*w

I

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A Flora! and. Fauna! Inventory of th
Par*: National de Marojejy,
Madagascar: With Reference to
ElevationaS Variation sf?w

Steven M. Goodman, ^dHo-

>i>t M, 2000

ri<i)I , ..;.Hi'ji

nl

J r-i. u :vj

LJSKUM OF NATURAL HISTORY

'

A Floral and Faunal Inventory
of the Pare National de
Marojejy, Madagascar

WWF

THE LIBRARY OF THE

SFP ! 1 2001

UNIVERSITY OF ILLINOIS
URBANA CHAMPAIGN

The Schlinger
Foundation

FIELDIANA

Zoology

NEW SERIES, NO. 97

A Floral and Faunal Inventory of the
Pare National de Marojejy,
Madagascar: With Reference to
Elevational Variation

Steven M. Goodman, Editor

Center for Environmental and Evolutionary Biology

Field Museum of Natural History

1400 South Lake Shore Drive

Chicago, Illinois 60605-2496

U.S.A.

World Wide Fund for Nature
B. P. 738

Antananarivo (101)
Madagascar

Accepted June 15, 1999
Published August 31, 2000
Publication 1510

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

2000 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

Contents

Preface vii

1. Description of the Pare National de Marojejy, Madagascar, and the 1996 Biological Invento-
ry of the Reserve 1

Steven M. Goodman
Appendix 1-1. Participants in the Project (Field and Laboratory) 17

2. Pteridophyte Diversity Patterns Along an Elevational Gradient in the Pare National de Maro-
jejy, Madagascar 19

France Rakotondrainibe

3. Structure and Floristic Composition of the Vegetation of the Pare National de Marojejy,
Madagascar 41

Nathalie Messmer, Pierre Jules Rakotomalaza, and Laurent Gautier

4. The First Beckeriella Williston from the Afrotropical Region: Two New Species from Mada-
gascar (Diptera: Ephydridae) 105

Wayne N. Mathis and David A. Grimaldi

5. The Tracheline Spider Genus Paccius (Araneae, Corinnidae) in the Pare National de Maroje-
jy, Madagascar 115

Norman I. Platnick

6. Fruit Flies as Ecological Indicators: Species Diversity and Abundance of Drosophilidae
(Diptera) Along an Altitudinal Transect in the Pare National de Marojejy, Madagascar 123

David A. Grimaldi, Eric L. Quinter, and Tarn Nguyen

7. Parasitic and Commensal Arthropods of Some Birds and Mammals of the Pare National de
Marojejy, Madagascar 1 37

Barry M. OConnor

8. Notes on a Small Collection of Fishes from the Pare National de Marojejy, Northeastern
Madagascar, with a Description of a New Species of the Endemic Genus Bedotia (Atherino-
morpha: Bedotiidae) 143

Melanie L. J. Stiassny and Ian J. Harrison

9. Herpetofaunal Species Diversity and Elevational Distribution Within the Pare National de
Marojejy, Madagascar 157

Achille P. Raselimanana, Christopher J. Raxworthy, and Ronald A. Nussbaum

10. Birds of the Pare National de Marojejy, Madagascar: With Reference to Elevational Distri-
bution 1 75

Steven M. Goodman, A. F. A. Hawkins, and Jean-Claude Razafimahaimodison

1 1 . Tenrecs (Lipotyphla: Tenrecidae) of the Pare National de Marojejy, Madagascar 20 1

Steven M. Goodman and Paulina D. Jenkins

1 2. Rodents of the Pare National de Marojejy, Madagascar 23 1

Michael D. Carleton and Steven M. Goodman

13. Rapid Census of Lemur Populations in the Pare National de Marojejy, Madagascar 265

Eleanor Sterling and Karen McFadden

Gazetteer of Localities Mentioned in the Text 275

Index to Scientific Names 279

Preface

In this volume we present the results of a brief
floral and faunal survey conducted in the Pare Na-
tional (PN) de Marojejy, northeastern Madagas-
car, between 4 October and 20 November 1996.
Since I began biotic inventories in Madagascar
several years ago, numerous reports and publica-
tions on the plants and animals of the Marojejy
Massif have fascinated me, in particular the vol-
ume produced by Henri Humbert in 1955 with the
tantalizing title Une merveille de la Nature a
Madagascar: Premiere exploration botanique du
massif du Marojejy et de ses satellites. Thus, we
were delighted to have the opportunity to visit this
mountain and continue the work started by nu-
merous other field scientists.

Our survey of the Marojejy Massif, organized
by the World Wide Fund for Nature (WWF),
Madagascar, was the fourth of five multidisciplin-
ary large-scale biological and elevational inven-
tories conducted on mountains in Madagascar
over the past 5 years. This volume is the fourth
inventory to be published in the Fieldiana: Zo-
ology series. The previous volumes were inven-
tories of the Reserve Naturelle Integrate (RNI)
d'Andringitra (now reclassified as a PN; Fieldi-
ana: Zoology, n.s., no. 85, 1996), the Reserve
Speciale (RS) d'Anjanaharibe-Sud {Fieldiana:
Zoology, n.s., no. 90, 1998), and the RNI
d'Andohahela (now reclassified as a PN; Fieldi-
ana: Zoology, n.s., no 94, 1999). Another volume
on the RS du Pic d'lvohibe and the corridor be-
tween this reserve and the PN d'Andringitra was
recently published in French in the series Re-
cherches pour le Developpement, Serie Sciences
Biologiques, no. 15 (1999).

Many people and institutions provided assis-
tance that made our inventory of the Marojejy
Massif possible. Our colleagues in the Malagasy
government, particularly the Direction des Eaux
et Forets, Association Nationale pour la Gestion
des Aires Protegees (ANGAP), and members of
the Commission Tripartite, provided the necessary
permits to carry out this work; we owe much to
their continued aid and collaboration. We are
deeply indebted to WWF staff members based in
Antananarivo and Andapa: Anjara Andriamanali-

na, Jean-Marc Garreau, Olivier Langrand (now of
WWF Gabon), Patricia Lehmann, Anjara Man-
antsara, Jean-Paul Paddack, Lantasoa Ramaroja-
ona, Gisele Ramaroson, and Malalarisoa Razafim-
pahanana. The assistance of the inhabitants of
Manantenina and Mandena is gratefully acknowl-
edged, in particular M. Zeze Bruno, Roland Chris-
tophe, and Bevao Jean Chrysostome. M. Razoki-
ny and M. Simona Arsene of Eaux et Forets, An-
dapa, helped with numerous aspects of this mis-
sion. We are also indebted in many ways to M.
Ledada Rachel Razafindravao, our highly talented
cook, who persevered under rather difficult con-
ditions. Movements up and down a mountain by
a research group and up to 40 porters can pose
numerous problems; however, the logistical aid
and competence of Mamy Ravokatra greatly fa-
cilitated this aspect of the mission.

The chapters in this volume have benefited
from the comments of numerous reviewers. Those
who waived anonymity are cited in the acknowl-
edgments to the chapters they reviewed. John
Weinstein, Field Museum of Natural History,
helped with the preparation of many of the pho-
tographs that appear in this volume. William Bur-
ger, Scientific Editor of Fieldiana, and Marjorie
Pannell, Managing Editor of the Field Museum
Press, continued to provide valuable assistance in
producing these volumes. Over the past 5 years,
they have seen nearly 3,000 manuscript pages as-
sociated with these Madagascar monographs pass
over their desks. For assistance with the transla-
tion of the abstracts into French, we thank Voni-
navoko Raminoarisoa, Bernardin Rasolonandra-
sana, and Lucienne Wilme.

The 1996 biological inventory of the PN de
Marojejy was made possible by grants from Kre-
ditanstalt fur Wiederaufbau (KfW) to WWF as
part of the Projet de Conservation et de Devel-
oppement Integres Marojejy/Anjanaharibe-Sud
and from the Center for Biodiversity and Conser-
vation of the American Museum of Natural His-
tory. KfW and the Schlinger Foundation provided
partial subsidy for the publication of this volume.

S. M. Goodman
3 June 1999
Chicago

Chapter 1

Description of the Pare National de Marojejy,
Madagascar, and the 1996 Biological Inventory of the
Reserve

Steven M. Goodman 1

The past 10 years have witnessed an impressive
expansion of scientific knowledge regarding Mada-
gascar resulting from a variety of sources sym-
posia focused on the island's history and biological
diversity (Ganzhorn et al., 1997; Goodman & Pat-
terson, 1997; Lourenco, 1996), long-term field stud-
ies on a variety of plants and animals at several
biological research stations (e.g., Ganzhorn & Sorg,
1996; Wright, in press); a wealth of behavioral, eco-
logical, and taxonomic studies on particular organ-
isms; rapid biological inventories of poorly known
forested areas on the island (Goodman, 1996, 1998,
1999; Goodman & Langrand, 1994; Langrand &
Goodman, 1997; Rakotondravony & Goodman,
1998; Ratsirarson & Goodman, 1998); and numer-
ous popular guides to its biotic splendors (Dransfield
& Beentje, 1995; Glaw & Vences, 1994; Langrand,
1990, 1995; Mittermeier et al., 1994; Morris &
Hawkins, 1998). This nearly exponential increase in
knowledge about Malagasy plants and animals has
enabled scientists to synthesize new ideas associated
with the biological patterns of the island to begin to
understand, for example, the effects of ecological
change during recent geological time. One of the
critical uses of the newly available information is to
strengthen conservation activities to safeguard what
remains of this truly remarkable biota.

Even with these advances in comprehending
the extant fauna and flora, major gaps remain in
basic knowledge of the natural history and distri-
bution of the great majority of Malagasy organ-
isms. Further, a remarkable number of taxa new
to science, including vertebrates, are being de-
scribed each year from Madagascar. Because of

1 Field Museum of Natural History, 1400 South Lake
Shore Drive, Chicago, IL 60605-2496, U.S.A.

the continued high levels of habitat destruction
(Whitmore, 1997) and the fact that very little of
the original forest cover is left, the coming decade
may be the last possible time to document the
remaining patterns of biotic diversity for numer-
ous sites on the island. Rapid biological invento-
ries of previously unstudied or poorly known for-
ested sites are critical to this effort.

This volume presents the results of a biological
inventory conducted in the Pare National (PN) de
Marojejy in northeastern Madagascar. The park lies
within the Province d'Antsiranana and between the
towns of Andapa and Sambava (Fig. 1-1). Fourteen
scientists participated in the field inventory, which
look place in the latter half of 1996.

In 1994 the eastern slopes of the Anjanaharibe-
Sud Massif was the site of an intensive biological
inventory (Goodman, 1998), part of a series of
inventories of mountains ranging along the com-
plete 13 latitudinal breadth that Madagascar en-
compasses. Data obtained on the massif, which
lies more to the interior of the island and on the
opposite side of the Andapa basin from the PN de
Marojejy (but at nearly the same latitude), provide
an excellent comparative base for examining pat-
terns of congruence in the distribution of many
different organisms along various latitudinal
clines. The authors of several chapters in this vol-
ume have compared the survey information from
the PN de Marojejy with the earlier data from the
RS d'Anjanaharibe-Sud.

This chapter briefly reviews the history of the
park, the organization of our 1 996 field season in
the park, geology, and climate. For further infor-
mation about the mountain readers are referred to
Humbert (1955), Guillaumet et al. (1975), Nicoll
and Langrand (1989), and Safford and Duckworth
(1990).

FIELDIANA: ZOOLOGY, N.S., NO. 97, AUGUST 31, 2000, PP. 1-286

ANTSIRANANA
PROVINCE

R.S. d'

Anjanaharibe Sud

MAHAJANGA
PROVINCE

Villages

Province limits
/V Coast

Fig. 1-1. Map of northeastern Madagascar showing the position of the Reserve Speciale d'Anjanaharibe-Sud and
the Pare National de Marojejy with respect to other sites, provincial limits, and towns in this part of the island.

Abbreviations Used in the Text

ANGAP Association Nationale pour la Gestion

des Aires Protegees
BM(NH) The Natural History Museum, London

(formerly British Museum [Natural

History])
CNRE Centre National de Recherche sur

l'Environnement, Antananarivo
CNRS Centre National de la Recherche Scien-

tifique, Antananarivo
dbh diameter at breast height

DEF Direction des Eaux et Forets

DRFP Direction des Ressources Forestieres et

Piscicoles
FMNH Field Museum of Natural History, Chi-
cago
FTM Foiben-Taosarintanin'i Madagasikara,

Antananarivo (Institut National de
Geodesie et Cartographie)

ha hectare

IRSM Institut de Recherche Scientifique de
Madagascar

KfW Kreditanstalt fur Wiederaufbau

MBG Missouri Botanical Garden, St. Louis

MNHN Museum National d'Histoire Naturelle,
Paris

MRAD Ministere de la Recherche Appliquee
au Developpement, Antananarivo

PBZT Pare Botanique et Zoologique de Tsim-
bazaza, Antananarivo

PN Pare National

RB Reserve de Biosphere

RCP La Recherche Cooperative sur Pro-

gramme No. 225, under the Centre Na-
tional de la Recherche Scientifique

FIELDIANA: ZOOLOGY

Fig. 1-2. Map of the approximate vegetational formations in the Pare National de Marojejy (modified after Razafy
Fara, 1998). The locations of the camps occupied during the 1996 biological inventory of the reserve are indicated.

RNI

RS

UADBA

UMMZ

USNM

WWF

Reserve Naturelle Integrate

Reserve Speciale

Universite d' Antananarivo Departe-

ment de Biologie Animale

University of Michigan Museum of

Zoology, Ann Arbor

National Museum of Natural History,

Washington, D.C. (formerly United

States National Museum)

World Wide Fund for Nature

The Marojejy Reserve

In early January 1952, after the exploration and
remarkable discoveries of Henri Humbert con-

cerning the flora on the Marojejy Massif, a por-
tion of this area was designated a reserve naturelle
(Nicoll & Langrand, 1989). This decree was fur-
ther modified on 1 June 1966 by another decree
(no. 66-242) naming 60,150 ha of the region as
the 12th Reserve Naturelle Integrate (RNI) de
Marojejy. Subsequently the status of this RNI was
changed to that of a park, Pare National (PN) no.
13, by decree 98-375, signed on 19 May 1998 and
published on 19 October 1998 (Gazetim-panjak-
an'ny, 1998), and the size of the park was reduced
to 60,050 ha. The principal reason for this change
in status was to provide opportunities for local
people to benefit from ecotourism and regulated
access to certain areas. Tourists have the right to
enter a national park after purchasing tickets, but

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

Table 1-1. Surface coverage in the PN de Marojejy, based on LANDSAT TM images.*

Vegetational type and surface coverage

Dense humid forest with closed canopy and multiple strata
Dense humid forest with open canopy and a single stratum
Secondary forest with patches of cleared areas
Herbaceous formations (including ferns)
Covered in clouds

Surface area

(ha)

Percent coverage

41,910

70.9

10,128

17.1

5,365

9.1

980

1.5

843

1.4

* Modified after Razafy Fara (1998).

protected areas with RNI status are only open to
scientists carrying special permits.

The PN de Marojejy comprises a wide range of
humid forest vegetational types, starting with low-
land forest at about 75 m and culminating in large
expanses above tree line of open ericoid bush and
exposed rock in the summital zone. The principal
peak is at 2132 m. The flora of the massif has
been studied by many botanists (Dorr, 1997;
Humbert, 1955), and a detailed checklist is cur-
rently in preparation (J. Miller, pers. comm.).
More detailed analyses of the vegetational zones
and the elevational stratification of the plant com-
munities are presented in Chapters 2 and 3. Also,
a number of works have been published on the
terrestrial fauna of the reserve (Duckworth et al.,
1995; Evans et al., 1992; Griveaud, 1960; Safford
& Duckworth, 1990).

A recurrent theme in this book is the excep-
tional biological diversity of the PN de Marojejy
in comparison with most of the reserves in the
protected areas system of Madagascar. This rich-
ness is a result of the park encompassing a broad
swath of elevational zones, starting from lowland
formations and ranging continuously to high
mountains.* The eastern foot of the Marojejy
Massif begins within a few tens of kilometers
from the Indian Ocean. In numerous other re-
serves on the island that include mountainous
zones, the lowest represented forest starts at a dis-
tinctly higher elevation (e.g. Anjanaharibe-Sud
and Andringitra). For many groups of land ver-
tebrates, such as birds and the herpetofauna, there
is a strong relationship between increasing ele-
vation and decreasing species richness. Simply
put, the lowland forests are the richest.

On the basis of patterns of vegetational cover-
age of the PN de Marojejy derived from LAND-
SAT TM images and ground-truthing (Razafy

* The terminology used for forest types on Madagas-
car is complicated. In Table 3-6 the various descriptive
names of these classifications are compared.

Fara, 1998), nearly 90% of the reserve consists of
dense, humid forest (Table 1-1). What is remark-
able about this figure is that a large percentage of
this habitat is forest below 800 m elevation (Fig.
1-2). We strongly suspect that the lowland forest
is important in explaining the reserve's exception-
ally high species richness. Few reserves in Mad-
agascar cover broad elevational ranges with this
high a percentage of lowland forest. An important
exception is the Tsaratanana Massif, which in-
cludes a variety of habitats, from 227 m to the
summit at 2876 m (Nicoll & Langrand, 1989).
The RNI de Tsaratanana has not been the site of
a multidisciplinary elevational inventory, and data
are not available to compare its richness with Ma-
rojejy's. However, once the site has been exten-
sively surveyed, it may show similarly high levels
of species richness.

Transect Sites

During the 1996 inventory of the Marojejy
Massif, coordinates for each site were determined
with the use of a geographical positioning system,
and the names for various localities were garnered
from maps (Foiben Taosarintanin'i Madagasikara
[FTM], 1974, 1977) and discussions with local
people. A gazetteer is presented at the end of this
volume (p. 275) for the localities mentioned in the
text.

Our inventory of the southeastern slopes of the
Marojejy Massif was conducted in an ascending
series of altitudinal transect zones, following the
classic route established by earlier explorers and
scientists who visited the site, such as L.-J. Ara-
gon. H. Humbert, P. Griveaud, the field team un-
der the direction of R. Paulian (Guillaumet et al..
1975), botanists from the Missouri Botanical Gar-
den (MBG), a group from the University of Lon-
don (Safford & Duckworth. 1990), and a number
of other researchers. We worked in five altitudinal

FIELDIANA: ZOOLOGY

Fig. 1-3. Lower portion of a largely undisturbed area of forest (at ca. 500 m) with a thick understory, relatively
dense lianas, and some remaining large trees. The vegetational types in the lowest elevational zone of the Marojejy
Massif were a mixture of secondary forest, dense areas of bamboo, and relatively intact lowland forest. (Photograph
by N. Messmer.)

zones (often referred to in this volume as transects
or transect zones) on the mountain that were gen-
erally delimited by a 75 m elevational band cen-
tered on each of the five camps. Elevations were
determined with the use of altimeters. The posi-

tion of each camp occupied during the survey is
indicated in Figure 1-2 (p. 3).

450 m (camp 1) 4-14 October 1996

Madagascar: Province d'Antsiranana, Pare Na-

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

Fig. 1-4. View of Ambatotsondrona rising to 1261 m, just across the Manantenina River valley from our camp
2 at 775 m and along the Ambavaomby River. The forest at the foot of the mountain has a discontinuous canopy
cover, presumably as a result of natural landslides, and secondary forest zones associated with human activities.
(Photograph by F. Rakotondrainibe.)

tional de Marojejy, along tributary of Manan-
tenina River, 8 km NW Manantenina,
1426.2'S, 4946.5'E.

775 m (camp 2) 14-24 October 1996

Madagascar: Province d'Antsiranana, Pare Na-
tional de Marojejy, along a tributary of the
Manantenina River, 10 km NW of Mananteni-
na, 1426.0'S, 4945.7'E.

1325 m (camp 3) 24 October-4 November 1996
Madagascar: Province d'Antsiranana, Pare Na-
tional de Marojejy, 1 1 km NW of Manantenina,
Antranohofa, 1426.2'S, 4944.5'E.

1625 m (camp 4) 4-13 November 1996
Madagascar: Province d'Antsiranana, Pare Na-
tional de Marojejy, 10.5 km NW of Mananten-
ina, along a tributary near the source of the An-
dranomifototra River, 1426.4'S, 4944.5'E.

1875 m (camp 5) 13-20 November 1996
Madagascar: Province d'Antsiranana, Pare Na-
tional de Marojejy, 1 1 km NW of Manantenina,

at the source of the Andranomifototra River,
1426.8'S, 4944.1'E.

Itinerary of the 1996 Expedition

From 3 to 6 September 1996, together with De-
sire Ravelonarivo, Gisele Ramaroson, and a local
guide, I conducted a quick reconnaissance of the
area along the Manantenina-summit trail up to
Antranohofa at 1250 m. Sites were chosen for our
first three camps. After we descended to the vil-
lage of Manantenina we met with the village el-
ders and local officials to explain our presence in
the area and discuss the plans for the inventory;
we also engaged local people to work with the
field group and established porter prices for the
various stages of the mission. I then returned to
Antananarivo by plane (via Sambava) to organize
other aspects of the mission.

On 30 September 1996 a vehicle driven by M.

FIELDIANA: ZOOLOGY

Fig. 1-5. View north, looking toward Ambatotsondrona (left center) and across the large areas of lowland forest
that remain in the PN de Marojejy. The photograph was taken from the trail above the upper limit of the elevational
transect centered at 775 m. The Manantenina River valley is just out of view at the bottom, (Photograph by S. M.
Goodman.)

Ravokatra, in the company of M. Ledada, left An-
tananarivo for Andapa loaded with field gear and
a good portion of the dried food provisions (ex-
cluding rice) for the mission. This journey, in a
four-wheel-drive vehicle, took 3 days. On 2 Oc-
tober 1 996 the group of researchers based in An-
tananarivo and those who had recently arrived
from other countries to join the survey team flew
from Antananarivo to Sambava, where they were
met by WWF vehicles and driven to Andapa. The
day of 3 October was spent in Andapa arranging
baggage and provisions, including 600 kg of rice
purchased locally, and paying courtesy visits to
various government offices. Research materials
and dried food provisions were separated accord-
ing to camp (transect zone), packed in labeled and
numbered rice sacks, and stored in a rat-proof, dry
building in Andapa.

Early on the morning of 4 October 1996 the
research group, baggage, and provisions for the
first camp were driven to the village of Manan-
tenina, where about 40 porters from this and
neighboring villages awaited our arrival. We then

walked 3-4 hours to a site at 450 m and estab-
lished our first camp.

Over the next 7 weeks the group, with the as-
sistance of porters from villages at the foot of the
mountain, changed study sites an additional four
times and then descended to Manantenina on 20
November 1996. Most of the group returned to
Andapa the same day, and on 22 November the
vehicle headed back to Antananarivo with the
specimens and research material; a number of the
researchers returned to Antananarivo by plane
(via Sambava).

The system for reprovisioning each camp was
as follows. One or two days before each sched-
uled displacement to the next camp, a WWF em-
ployee in Andapa purchased fresh produce in the
local market, and these goods, along with the
dried provisions and research materials stocked in
Andapa for the next camp, were transported to
Manantenina. Prearranged rendezvous were or-
ganized with local porters to carry the material to
the site we were preparing to leave. Once the por-
ters had reached the research group, the baggage,

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

Fig. 1-6. View toward the northeast from an open area at about 1250 m and below camp 3 at Antranohofa. The
peak in the center is Ambatotsondrona. a prominent feature opposite camp 2 at 775 m. Note the growth of bamboo
in the valley bottom in the foreground and changes in vegetational cover along exposed ridges. (Photograph by S.
M. Goodman.)

which included the new goods, goods from the
existing camp, and the porters" food provisions,
was reorganized and redistributed. This process
took from a few minutes to several hours, and
after it was completed the porters and field-work-
ers climbed to the next site. Specimens from each
site were temporarily stored in the abandoned
camp. After each displacement, the porters on
their way back down the mountain carried the
specimens to Manantenina. where a WWF vehicle
was waiting to transfer the specimens to the stor-
age building in Andapa.

After the third camp the distances between the
lowlands and our transect zones were too long to
complete the round-trip during 1 day. After our
displacement to camp 4 the porters spent the night
in camp 2. and after we moved to camp 5 the
porters passed the night at camp 3. On both oc-
casions these camps had to be stocked with tarps
for shelter, cooking pots, and provisions for the
porters. Between 32 and 42 porters were needed
for each displacement, with the exception of the
final descent, which did not involve any provi-

sions and took about 25 people, not including the
research group.

Compared with our study group's other eleva-
tional transects of mountainous regions in Mada-
gascar, the survey of the Marojejy Massif was rel-
atively simple from the point of view of logistics.
The Manantenina-summit trail was established
decades ago and has remained open owing to fre-
quent use. People in the local villages are accus-
tomed to working as porters and guides for sci-
entific groups, distances between sites can be cov-
ered in a few hours, and the climb to the summital
zone is not particularly steep.

During the field trip to the PN de Marojejy (4
October-20 November 1996) members of the sci-
entific crew included S. M. Goodman (birds and
small mammals), A. F. A. Hawkins (birds camps

1 and 2 only), J. Legrand (insects camps 1 and

2 only), N. Messmer (plants), T. Nguyen (in-
sects camps 1 and 2 only), E. Quinter (insects),
P. J. Rakotomalaza (plants), F. Rakotondrainibe
(ferns). B. Randriamampionina (ferns), D. Ran-
driamasimanana (insects camps 1 and 2 only),

FTELDIANA: ZOOLOGY

Fig. 1-7. Starting at about 1250 m, there are numerous areas of exposed rock or very shallow soils supporting
bamboos, ferns, or small geophytic plants. This photograph was taken at about 1275 m and below camp 3 on the
main trail leading down to camp 2. The view is toward the south. (Photograph by S. M. Goodman.)

A. Raselimanana (reptiles and amphibians), D.
Ravelonarivo (plants), M. Ravokatra (birds and
logistics), and J.-C. Razafimahaimodison (birds).
The group also included two to three local peo-
ple from the villages of Mandena and Mananten-
ina, a cook, and visitors from the WWF project

in Andapa. The maximum number of people we
had simultaneously in any one camp was about
22, not including porters. The names and address-
es of all field and laboratory researchers who took
part in this project are given at the end of this
chapter (Appendix 1-1).

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

iM$

Fig. 1-8. View of exposed ridge at about 1600 m showing largely sclerophyllous vegetation. These slopes are
regularly exposed to strong winds, heavy fog, and mist. (Photograph by N. Messmer.)

Logistics, Trail Systems, and
Descriptions of the Various Study
Zones

Our trajectory up the mountain followed the
Manantenina-summit trail, and the first three
camps were at sites already established along this
path by researchers who had previously visited
this slope of the massif. The vast majority of re-
cent scientists who visited the massif used the An-
tranohofa site at 1325 m (our camp 3) as their
base camp for work in the summital zone and
climbed up and down each day. At a stiff pace it
is possible to ascend from Antranohofa to the
summital zone in about 4 hours. However, for our
purposes this was not adequate, and we decided
to establish two other camps between Antrano-
hofa and the summital zone.

Chapters 2 and 3 of this volume present de-
tailed descriptions of the pteridophyte and angio-
sperm plant communities occurring within each
elevational zone. Here I restrict my comments to
aspects of access, trails, and the general condition
of each transect zone.

450 m

Our study area was close to the main valley of
the Manantenina River and in an area of disturbed
forest, at a site known locally as Ampanasanko-
lana (Ampanasama Ankolony) or Bedinta. The
camp was near the junction of two tributaries of
the Manantenina River and about 4 hours' walk
from Manantenina. The names of these tributaries
were not consistent between local informants but
are probably the Ambinantelo and Ampanasaton-
gotra Rivers.

The habitats within the transect varied from
heavily disturbed sites with low secondary growth
(savoka) to zones of dense bamboo mixed with
Aframomum (Zingiberaceae) to slightly disturbed
lowland humid forest habitat (Fig. 1-3). Portions
of this area had been cleared in the not too distant
past for agricultural use, particularly coffee plan-
tations (Humbert, 1955) and slash-and-burn prac-
tices (tavy). Most of the zones of secondary
growth were regenerating tavy plots, as were
probably also some of the nearly monospecific ar-
eas of bamboo. An extensive trail system existed

10

FIELDIANA: ZOOLOGY

Fig. 1-9. Camp 5 at 1875 m, above the forest line. The vegetation immediately around the camp was a mixture
of ericoid bush, dwarf palms, and herbaceous plants. The ground in low-lying areas was moist. The source of the
Andranomifototra River is in this region. (Photograph by F. Rakotondrainibe.)

in this portion of the park and is regularly used
by local people for collecting forest plants (me-
dicinal and construction uses), hunting and fish-
ing, and access to illegal tavys well hidden in var-
ious portions of the reserve. Photographs pub-
lished in Humbert's monograph (1955, pi. 24, figs.
47 and 48) show the hills above Mandena and
toward Mt. Behondroko as already a patchwork
of forest and agricultural plots. In 1996 the level
of habitat degradation was much more severe, and
little forest remained on these hills.

775 m

This site was next to the Andampimbazaha
Cascade, a waterfalls along the Ambavaomby
River about 100-150 m elevation above where it
plunges into the Manantenina River. Across the
main river valley is the distinct rock outcrop
known as Ambatotsondrona (Fig. 1-4). The walk
to the 775 m camp from the 450 m site takes
about 1 hour and is along the trail leading to the
summit.

In general, the forest below the 775 m camp
site showed signs of human disturbance, mostly
plant wood extraction for building materials,
while above the camp the forest seemed largely
intact. It was in this zone that the magnificent Ma-
rojejya palms, a genus discovered during Hum-
bert's explorations on the Marojejy Massif, were
clearly in evidence. Small trails existed through-
out the upper portion of this transect zone, and
these paths often ended in a small and well-hidden
tavy, most of which had been recently abandoned,
or at a site where a tree had been removed. Thus,
it was clear that until very recently, local people
harvesting forest products from the park had tried
to conduct their activities off the beaten track. The
Manantenina Valley and the low hills to the north
still hold considerable areas of the original low-
land and montane forest (Fig. 1-5).

1250 m

This site, known as Antranohofa, was within 3
hours' walking distance of camp 2 and along the

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

11

I

Fig. 1-10. The habitats occurring in the zone above camp 5 at 1875 and the summit varied from low ericoid
bush, often with stands of bamboo and palms in the valley bottoms, to areas of low-growing herbaceous and woody
vegetation, and finally bare exposed rock. The large dome at the top of the photograph is the summit of Marojejy.
at 2132 m. (Photograph by F. Rakotondrainibe.)

trail leading to the summit. We designated the
camp at 1325 m as the upper limit of the transect
zone, with the midpoint 75 m below, at 1250 m.
This locality is probably the Andilana of Humbert
(1955) and the Andasy II of Griveaud (1960). Just

next to the campsite is a small tributary that has
potable water most of the year, except at the end
of the dry season. In early September, during the
reconnaissance mission, this source was nearly
dry. However, when we arrived to occupy the site

12

FIELDIANA: ZOOLOGY

Motion O.R.9.T.O.M.XU-Vg5fl Rc'fleryg N'U .
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Fig. 1-11. Messages left in a container cached in a small cairn at the principal summit of the Marojejy Massif
at 2132 m. Top, A note left by P. Griveaud concerning his December 1958 trip to the mountain describes the members
of the group and the sites they visited. Much of the information collected by Griveaud during this trip focused on
insects, but he also published some information on the birds and mammals occurring on the massif (Griveaud, 1960).
When he visited the summit in 1958 he found numerous slips of paper left behind by earlier visitors to the site. Most
slips had badly deteriorated. Griveaud recopied certain messages of historical interest and added the fresher slip of
paper to the cairn bottle (bottom). Among the notes in his distinctive handwriting is information on the 1937 visit
of Aragon, the first person to reach the summit, and records of passage of such notable individuals as Professor H.
Humbert. The Griveaud recapitulation is also in poor shape, and we reproduce it here to continue the archival tradition.

in late October there was a considerable amount
of water. In addition, there was a permanent
stream within 100 m by foot below the camp.
There were few signs of human activity in the
Antranohofa transect, with the exception of an oc-
casional cut tree of Evodia sp. (bilahy), the bark
of which is used to make an alcoholic beverage
known as betsa-betsa.

About 400 m walking distance below the An-
tranohofa camp, on the trail leading back to camp
2, there is a precipitous drop in the topography,
from which one can look down into the Manan-
tenina River valley. From this lookout point it is
also possible to see Ambatotsondrona, the massif
just across the Manantenina Valley from our camp
2 (Fig. 1-6). In this area of the trail there were

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

13

Table 1-2. Summary of minimum and maximum temperatures and precipitation during the 1996 expedition to
PN de Marojejy.

Periods of measurement
within each transect

Temperature CO*

Minimum

Maximum

Rainfall (mm)f

450 m

4-14 October

750 m

15-23 October

1325 m

24 October-3 November

1550 m

4-13 November

1875 m

14-20 November

10, 13.0-17.0
14.7 1.10

10, 13.0-18.0
15.9 1.58

9, 10.0-13.0
11.4 0.83
9, 9.0-13.0
11.0 1.40
7, 8.0-13.0
9.9 1.64

10, 17.0-23.0

20.1 1.87
10. 19.0-27.0

23.2 2.14
9, 19.0-23.0

21.3 1.56
9. 15.0-18.0

16.5 1.17
7. 18.0-23.0
21.0 1.69

8, 2.0-40.0
10.6 12.68

7. 1.0-27.5
12.2 10.76

5, 1.0-23.0
8.0 7.8

6, 0.5-45.0
11.0 15.84

5. 0.5-1.0
0.6 0.2

Data are presented as number of records, range, mean, and standard deviation.

Data are presented as number of days with rain, range, mean, and standard deviation.

numerous sites with exposed rock or very shallow
soils supporting bamboo, ferns, or small geophy-
tic plants (Fig. 1-7).

1550 m

Our camp 4 was about 1% hours' walk from
camp 3. The site was at 1550 m, which marked
the lower end of the transect (1550-1700 m, cen-
tered at 1625 m). The camp was along the An-
dranomifototra River and accessible by a newly
cut path, about 1 km from the main summit trail.
The camp was within an extensive area of mon-
tane and mossy forest. The only signs of human
activity that we found in this area were a few trees
of bilahy that had been cut and their bark re-
moved.

This zone encompassed the ecotones between
montane and sclerophyllous forest, particularly on
ridges that were exposed to wind (Fig. 1-8). In
the protected valleys the vegetational structure
had more of a montane forest aspect.

1875 m

The site of our camp 5 was above the tree line
on the flat portion of a ridge overlooking the
source of the Andranomifototra River (Fig. 1-9).
The camp was about 300 m from the main summit
trail and a 1.5-2 hours' climb from camp 4. The
site afforded easy access to the summital zone,
the open ericoid bush, and the upper limit of the
sclerophyllous forest. On clear days we had spec-
tacular views of the Lokoho and Manantenina riv-

er systems, the Indian Ocean, and at night the
lights of Sambava. The walk from the camp to the
summit took about an hour. The habitats in this
region varied from low ericoid bush, often with
stands of bamboo and palms in the valley bot-
toms, to areas with low-growing herbaceous and
woody vegetation, and finally bare exposed rock
(Fig. 1-10).

As mentioned earlier, numerous missions to
Marojejy had reached the summit of the massif.
It is customary among mountain enthusiasts on
Madagascar to place notes recording their passage
in a cairn at the summits of the major peaks. At
the summit of Marojejy we found an old jar with
messages left by groups that had successfully
reached this point. Several of the older messages
or those written on poor-quality paper were in
poor condition, and some were already illegible.
Messages that we considered important from a
historical perspective were carried back to An-
dapa by Jean-Marc Garreau, photocopied, re-
turned to the site, and placed in a new, hermeti-
cally sealed plastic container. Two of these mes-
sages had been written by R Griveaud in his dis-
tinctive penmanship, one recounting his ascent to
the summit in 1958 and the other recapitulating
portions of messages in poor condition found at
the time of his passage (Fig. 1-11).

Geology

The Marojejy Massif covers an area approxi-
mately 32 km from east to west and 22 km from
north to south. The massif is lithologically com-

14

FIELDIANA: ZOOLOGY

co r-

in

ON ON

NO
CM

o

CO CM

O 00
CO

On m

CM 00

ON Tj"

CM

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00

o< d

CM CM

ON NO

CM

i/~i no

TT

00 ON

ON t

NO

CM

so q

(N NO

00 NO

r^ oo

o

SO

CM

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"st oo

co

no r-'

"

O CM

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00 CM

CM

00 CO

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TT ON

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io r-t

r-

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in on

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^r

so

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00 CM
CM

CM

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sO

ON NO

sO

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^t CM

CO

CO CM
U0 CM

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00

r~ co

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CO CM

*s

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O
en NO

>>ON

C3

ON Tf

CM

Tj- CM

d o<

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1 Eco^ I E

2 3 no E 3

cs .a a

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plex, with portions being granitic, granite-dioritic,
malagachitic, and a section toward the summit
area composed of gneiss. On the basis of infor-
mation presented by Humbert (1955), Karche and
Petit (1969), and Petit (1971), I will briefly de-
scribe the geology of the massif along the Man-
antenina summit slope. Above the main Manan-
tenina River valley the first major cascade occurs
above 700 m (in the vicinity of our camp 2) and
the rock is generally a form of gneiss interstrati-
fied with charnockitic, syenitic, and migmatitic
rock. Between about 700 m and 1200 m the for-
mation is largely charnockite and syenite. From
about 1200 m the rock, which rises to form the
summit of the mountain, is principally granite,
with intrusions of white and rose quartz.

Meteorology

One of the difficulties in interpreting weather
patterns in the PN de Marojejy is the lack of data
from the immediate environs of the massif: the
nearest weather stations are along the coast, at
Sambava (about 50 km direct distance), and fur-
ther inland, at Andapa (about 50 km direct dis-
tance) (Fig. 1-1). The mean annual precipitation
is 2,296 mm at Sambava and 1 ,883 mm at Andapa
(Donque, 1975). Rainfall is probably higher on
the Marojejy Massif, and there is certainly ele-
vational variation on the mountain with respect to
precipitation and temperature. To establish a
weather profile for Marojejy, we measured mini-
mum and maximum daily temperatures (C) and
amount of precipitation every 24-hour period (Ta-
ble 1-2).

As expected, daily minimum temperatures were
generally lower with increasing elevation. An ex-
ception was the mean minimum temperature at
750 m (15.9C), which was warmer than at 450
m (14.7C). One explanation for this observation
is that the 750 m campsite was in an exposed area
with considerable amounts of open dark-colored
rock that absorbed and retained a greater amount
of solar energy. Another possibility is that the val-
ley in which our 450 m camp was located acts as
a cold air funnel.

In contrast to minimum temperatures, daily
maximum temperatures across the altitudinal gra-
dient deviated substantially from a simple model
of decreasing values with increasing elevation.
The 750 m camp had the highest mean daily tem-
perature, perhaps because of the exposed nature

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

15

of the site. The 1330 m site had a slightly higher
mean daily maximum than the site at 450 m. The
camp at 1875 m was in an open area above the
tree-line and exposed to direct solar radiation,
which accounts for the high daily temperatures
relative to the 1625 m site.

Levels of rainfall showed no clear pattern in
relation to the elevational sequence of our study
sites. To a large extent the amount of rainfall was
equal across the four lower zones within the for-
est. The main differences between the five sites
occurred at the 450 m and 1675 m camps, where
several heavy showers (40-45 mm) swelled the
average daily rainfall figure. Although the rainy
season in this part of Madagascar normally begins
in earnest by December, there was no clear pattern
of an increase in number of days with rain per
transect zone during our inventory of Marojejy.
At Sambava, the driest month over the course of
the year on average is October (Petit, 1971; Table
1-3, this chapter). Eighty percent of the days we
were present at 450 m had rain, 70% at 750 m,
55% at 1330 m, 66% at 1675 m, and 71% at 1875
m. The most unusual rainfall pattern was seen at
the 1875 m camp. During the 7 days we were
present at the site, the maximum rainfall on any
given day was 1.0 mm, and in general most pre-
cipitation occurred as dew and heavy morning
mist.

Literature Cited

Donque, G. 1975. Contribution geographique a l'etude
du climat de Madagascar. Nouvelle Imprimerie des
Arts Graphiques, Antananarivo, vii + 478 pp.

Dorr, L. J. 1997. Plant collectors in Madagascar and
the Comoro Islands. Royal Botanic Gardens, Kew,
xlvi + 524.

Dransfield, J., and H. Beentje. 1995. The palms of
Madagascar. Royal Botanic Gardens, Kew, xii + 475
pp.

Duckworth, J. W., M. I. Evans, A. F. A. Hawkins, R.
J. S afford, and R. J. Wilkinson. 1995. The lemurs
of Marojejy Strict Nature Reserve, Madagascar: A sta-
tus overview with notes on ecology and threats. In-
ternational Journal of Primatology, 16: 545-559.

Evans, M. I., J. W. Duckworth, A. E A. Hawkins, R.
J. Safford, B. C. Sheldon, and R. J. Wilkinson.
1992. Key bird species of Marojejy Strict Nature Re-
serve, Madagascar. Bird Conservation International,
2: 201-220.

Foiben Taosarintanin'i Madagasikara. 1974. Ma-
roambihy, 1:100,000, feuille XY-36. Foiben Taosar-
intanin'i Madagasikara, Antananarivo.

. 1977. Andapa, 1:100,000, feuille W-36. Foiben

Taosarintanin'i Madagasikara, Antananarivo.

Ganzhorn, J. U., B. Rakotosamimanana, L. Hannah,
J. Hough, L. Iyer, S. Olivieri, S. Rajaobelina, C.
Rodstrom, and G. Tilkin. 1997. Priorities for biodi-
versity conservation in Madagascar. Primate Report,
48-1: 1-81.

Ganzhorn, J. U., and J. -P. Sorg, eds. 1996. Ecology
and economy of a tropical dry forest in Madagascar.
Primate Report, Special Issue, 46-1: 1-382.

Gazetim-panjakan'ny. 1998. Decret no. 98-375 portant
changement de statut de la Reserve naturelle integrale
no. 12 de Marojejy en Pare national no. 13. Gazetim-
panjakan'ny Repoblikan'i Madagasikara, 19 Oktobra
1998, pp 3137-3144.

Glaw, F, and M. Vences. 1994. A fieldguide to the
amphibians and reptiles of Madagascar. Zoologisches
Forschungsinstitut und Museum Koenig, Bonn, 480 pp.

Goodman, S. M., ed. 1996. A floral and faunal inventory
of the eastern slopes of the Reserve Naturelle Integra-
le d'Andringitra, Madagascar: With reference to ele-
vational variation. Fieldiana: Zoology, n.s., 85: 1-319.

., ed. 1998. A floral and faunal inventory of the

Reserve Speciale d'Anjanaharibe-Sud, Madagascar:
With reference to elevational variation. Fieldiana: Zo-
ology, n. s., 90: 1-246.

-., ed. 1999. A floral and faunal inventory of the

Reserve Naturelle Integrale d'Andohahela, Madagas-
car: with reference to elevational variation. Fieldiana:
Zoology, n. s., 94: 1-297.

Goodman, S. M., and O. Langrand, eds. 1994. Inven-
taire biologique: Foret de Zombitse. Recherches Pour
le Developpement, Serie Sciences Biologiques, Centre
d'Information et de Documentation Scientifique et
Technique, Antananarivo, No. Special, 106 pp.

Goodman, S. M., and B. D. Patterson. 1997. Natural
change and human impact in Madagascar. Smithson-
ian Institution Press, Washington, D.C., xiii + 432 pp.

Griveaud, P. 1960. Une mission entomologique au Ma-
rojejy. Naturaliste Malgache, 12: 43-55.

Guillaumet, J.-L., J.-M. Betsch, C. Blanc, P. Morat,
and A. Peyrieras (under the direction of R. Paulian).
1975. Etude des ecosystemes montagnards dans la re-
gion malgache. III. Le Marojezy. IV. LTtremo et
l'lbity. Geomorphologie, climatologie, faune et flore
(Campagne RCP 225, 1972-1973). Bulletin du Mu-
seum National d'Histoire Naturelle, 3e serie, Ecologie
generate, no. 309, 25: 29-67.

Humbert, H. 1955. Une merveille de la Nature a Mad-
agascar. Premiere exploration botanique du massif du
Marojejy et de ses satellites. Memoires de l'lnstitut
Scientifique de Madagascar, serie b, 6: 1-210.

Karche, J. P., and M. Petit. 1969. Grand traits geo-
morphologiques du Marojejy central. Semaine Geo-
logique (Antananarivo), pp. 105-109.

Langrand, O. 1990. Guide to the birds of Madagascar.
Yale University Press, New Haven, xi + 364 pp.

. 1995. Guide des oiseaux de Madagascar. De-

lachaux et Niestle, Lausanne, 415 pp.

Langrand, O., and S. M. Goodman, eds. 1997. Inven-
taire biologique: Foret de Vohibasia et d'lsoky-Vohi-
mena. Recherches Pour le Developpement, Serie Sci-

16

FIELDIANA: ZOOLOGY

ences Biologiques, Centre d' Information et de Docu-
mentation Scientifique et Technique, Antananarivo,
no. 12, 197 pp.
Lourenco, W. R., ed. 1996. Biogeographie de Mada-
gascar. ORSTOM Editions, Paris, X + 588 pp.

MlTTERMEIER, R. A., I. TATTERSALL, W. R. KONSTANT, D.

M. Meyers, and R. B. Mast. 1994. Lemurs of Mad-
agascar. Conservation International, Washington,
D.C., 356 pp.

Morris, P. A. J., and A. F. A. Hawkins. 1998. The birds
of Madagascar: A photographic guide. Pica Press,
East Sussex, xi + 316 pp.

Nicoll, M. E., and O. Langrand. 1989. Madagascar:
Revue de la conservation et des aires protegees. World
Wide Fund for Nature, Gland, xvii + 374 pp.

Petit, M. 1971. Contribution a l'etude morphologique
des reliefs granitiques a Madagascar. LTmprimerie
Centrale, Antananarivo, 307 pp.

Rakotondravony, D., and S. M. Goodman, eds. 1998.
Inventaire biologique: Foret d'Andranomay, Anjozo-
robe. Recherches Pour le Developpement, Serie Sci-
ences Biologiques, Centre d'lnformation et de Docu-
mentation Scientifique et Technique, Antananarivo,
no. 13, 110 pp.

Ratsirarson, J., and S. M. Goodman, eds. 1998. In-
ventaire biologique de la Foret Littorale de Tampolo
(Fenoarivo Atsinanana). Recherches Pour le Devel-
oppement, Serie Sciences Biologiques, Centre
d'lnformation et de Documentation Scientifique et
Technique, Antananarivo, no. 14, 261 pp.

Razafy Fara, F. L. 1998. Rapport sur la typologie de
la vegetation et des occupations de sol dans le Pare
National de Marojejy et du corridor forestier. Depar-
tement des Eaux et Forets, Antananarivo, 18 pp.

Safford, R. J., and J. W Duckworth, eds. 1990. A
wildlife survey of Marojejy Reserve, Madagascar.
ICBP Study Report 40. International Council for Bird
Preservation, Cambridge, England.

Whitmore, T C. 1997. Tropical forest disturbance, dis-
appearance, and species loss, pp. 3-12. In Laurence,
W. E, and R. O. Bierregaard, Jr., eds., Tropical forest
remnants: Ecology, management, and conservation of
fragmented communities. University of Chicago
Press, Chicago, xv + 616 pp.

Wright, P. C, ed. In press. Biodiversity science and
village economics: Madagascar's Ranomafana Nation-
al Park. Covelo, Island Press.

Appendix 1-1

Participants in the Project
(Field and Laboratory)

A total of 24 scientists and field-workers from
five different countries were involved in this mul-
tidisciplinary study. This number included field
participants listed in the previous section as well
as researchers responsible for some of the labo-
ratory studies. The addresses of all scientific par-
ticipants are given below.

Carleton, M. D., Division of Mammals, National Mu-
seum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Gautier, L., Conservatoire et Jardin Botaniques de la
Ville de Geneve, Case Postale 60, CH-1292, Cham-
besy/Geneve, Switzerland.

Goodman, S. M., Field Museum of Natural History,
1400 South Lake Shore Drive, Chicago, Illinois
60605-2416, U.S.A., and World Wide Fund for Na-
ture, Aires Protegees, B.P 738, Antananarivo (101),
Madagascar.

Grimaldi, D. A., Division of Invertebrate Zoology,
American Museum of Natural History, Central Park
West at 79th Street, New York, New York 10024-
5192, U.S.A.

Harrison, I. J., Department of Ichthyology, American
Museum of Natural History, Central Park West at 79th
Street, New York, New York, 10024-5192, U.S.A.

Hawkins, A. F. A., B.P. 8511, Antananarivo (101), Mad-
agascar.

Jenkins, P. D., Mammal Group, The Natural History
Museum, Cromwell Road, London SW7 5BD, United
Kingdom.

Legrand, J., Laboratoire d'Entomologie, Museum Na-
tional d'Histoire Naturelle, 45, rue Buffon, 75005 Par-
is, France.

Mathis, W N., Department of Entomology, NHB 169,
National Museum of Natural History, Smithsonian In-
stitution, Washington, D.C. 20560, U.S.A.

Messmer, N., Conservatoire et Jardin Botaniques de la
Ville de Geneve, Case Postale 60, CH-1292 Cham-
besy/Geneve, Switzerland.

Nguyen, T, Division of Invertebrate Zoology, American
Museum of Natural History, Central Park West at 79th
Street, New York, New York 10024-5192, U.S.A.

Nussbaum, R. A., Museum of Zoology, University of
Michigan, Ann Arbor, Michigan 48109-1079, U.S.A.

OConnor, B. M., Museum of Zoology, University of
Michigan, Ann Arbor, Michigan 48109-1079, U.S.A.

Platnick, N. I., Division of Invertebrate Zoology, Amer-
ican Museum of Natural History, Central Park West
at 79th Street, New York, New York 10024-5192,
U.S.A.

Quinter, E., Division of Invertebrate Zoology, American
Museum of Natural History, Central Park West at 79th
Street, New York, New York 10024-5192, U.S.A.

Rakotomalaza, P. J., Missouri Botanical Garden, B.P.
3391, Antananarivo (101), Madagascar.

Rakotondrainibe, E, Ecole Pratique des Hautes Etudes,
16, rue Buffon, 75005 Paris, France.

Randriamampionina, B., World Wide Fund for Nature,
B.P. 42, Tolagnaro (614), Madagascar.

Randriamasimanana, D., LRSAE/ORSTOM, B.P. 434,
Antananarivo (101), Madagascar.

Raselimanana, A., World Wide Fund for Nature, B.P.
738, Antananarivo (101), Madagascar.

Ravelonarivo, D., World Wide Fund for Nature-Andapa,
B.P. 34, Sambava (208), Madagascar.

Ravokatra, M., World Wide Fund for Nature, B.P. 738,
Antananarivo (101), Madagascar.

GOODMAN: DESCRIPTION OF PARC NATIONAL DE MAROJEJY

17

Rax worthy. C. J.. Division of Herpetology. American Stiassny. M. L. J.. Department of Ichthyology. Amer-
Museum of Natural History. Central Park West at 79th ican Museum of Natural Historv. Central Park West

Street. New York. New York 10024-5192. U.S.A. at 79th Street. New York. New York 10024-5192.

Razafimahaimodison. J.-C. Department of Ornithology. U.S.A.

American Museum of Natural Historv. Central Park
West at 79th Street. New York. New York. 10024-
5192. U.S.A.

18 FIELDIANA: ZOOLOGY

Chapter 2

Pteridophyte Diversity Patterns Along an Elevational
Gradient in the Pare National de Marojejy,
Madagascar

France Rakotondrainibe 1

Abstract

A study of the pteridophytes of the Pare National de Marojejy was conducted as part of an
ongoing biological inventory program of protected areas in Madagascar, operating since 1993.
A total of 239 species or infraspecific taxa in 70 genera were recorded. Of that total, 105
(43.9%) are endemic to Madagascar, and 6 are known only from the park. Its high level of
floristic richness may be explained in large part by several observations: the Marojejy reserve
lies in the wettest part of the island; its complex geomorphology generates a great number of
biotopes; moist evergreen forest covers a wide altitudinal range, from 400 to 2000 m; and its
former status as a strict nature reserve has been retained until mid- 1998. Compared to the
pteridophyte floras of other massifs studied in Madagascar (Anjanaharibe-Sud and Andohahela),
that of Marojejy has a relatively large number of species in the genera Lindsaea and Grammitis,
and lower richness in Asplenium, although the latter is still the dominant genus as elsewhere
in the island. Floristic data gathered on 194 species and varieties observed in 18 plots covering
a total area of 14,100 m 2 were analyzed using two complementary numerical methods: corre-
spondence analysis and hierarchical ascending classification. Four floristic groups were distin-
guished between 400 and 2000 m elevation on the eastern slope of Marojejy, each of which
was characterized regarding several features, including generic and specific composition, av-
erage density of the species present, "exclusively characteristic" and "preferentially charac-
teristic" species, and the range and percentages of growth forms present. The altitudinal zones
occupied by these floristic groups were compared to those of the vegetation types previously
described on the massif. Floristic similarity coefficients were calculated for the pteridophytes
occurring in each altitudinal zone on Marojejy for comparison with those in the corresponding
zones on Anjanaharibe-Sud ca. 35-40 km farther inland; the results show that the pteridophyte
flora of Marojejy is most distinctive at middle altitudes, between 1250 and 1350 m.

Resume

L'etude des Pteridophytes du Pare National de Marojejy a ete realisee dans le cadre des
campagnes d'inventaire biologique menees depuis 1993 dans les aires protegees de Madagascar.
Un total de 239 especes ou varietes reparties en 70 genres a ete recense. Parmi elles, 105 soit
43,9% sont endemiques de Madagascar et 6 endemiques du pare. La position geographique du
massif du Marojejy dans la region la plus arrosee de l'lle, la geomorphologie complexe du
massif qui abrite de nombreux biotopes, sa couverture forestiere continue entre 400 et 2000 m

1 Laboratoire de Biologie et Evolution des Plantes Vasculaires, Ecole Pratique des Hautes Etudes, 16, rue Buffon,
75005 Paris, France.

RAKOTONDRAINIBE: PTERIDOPHYTES 19

d'altitude et le statut de reserve integrate qui lui a ete attribue entre 1952 et 1998, expliquent
en grande partie cette richesse floristique elevee. Les traits caracteristiques de cette flore par
rapport a celle d'autres massifs malgaches (Anjanaharibe-Sud et Andohahela) sont la richesse
relative en taxons appartenant aux genres Lindsaea et Grammitis et une moindre diversite en
Asplenium qui reste cependant le genre dominant comme sur l'ensemble de l'lle. Les donnees
floristiques relevees sur 18 parcelles totalisant 14.100 m 2 et concernant 194 especes ou varietes
sont traitees par deux types d'analyses multivariables complementaires: une classification as-
cendante hierarchique et une analyse factorielle des correspondances. Quatre groupements flor-
istiques sont ainsi mis en evidence. Pour chacun d'eux, nous precisons: la composition gene-
rique et specifique, la densite moyenne des especes, les especes caracteristiques exclusives et
preferentielles, le spectre des types biologiques. La position de ces groupements par rapport a
l'etagement connu des formations vegetales du pare est discutee. La comparaison des coeffi-
cients de similitude floristique, etage par etage, des quatre groupements de Pteridophytes du
Marojejy avec ceux du massif de l'Anjanaharibe-Sud distant seulement de 35-40 km montre
que la specificite de la flore du Marojejy est la plus grande au niveau des altitudes moyennes,
entre 1250 et 1350 m.

Introduction

Of all the large mountain massifs in Madagas-
car, Marojejy was the last to be discovered but
has the best known flora (Des Abbayes, 1961;
Guillaumet et al., 1975; Humbert, 1955; Koechlin
et al., 1974). Humbert (1955) was the first botanist
to collect in the region. Between 1948 and 1951
he traveled throughout the principal massif of Ma-
rojejy as well as the massifs of Beondroka and
Mainborondro, located slightly to the east, Am-
batosoratra, to the south, Anjenabe and Betso-
manga, to the north, and several surrounding val-
leys, including the Lokoho, Antsahabe, Manan-
tenina, and Androranga. The pteridophyte speci-
mens that Humbert took back to the Museum
National d'Histoire Naturelle (MNHN) in Paris
were studied by Tardieu-Blot, who published (in
Humbert, 1955) a preliminary list of 107 species
and varieties of pteridophytes, including descrip-
tions of 1 1 new species, 8 of which were consid-
ered to be locally endemic: Asplenium andapense
Tardieu, A. marojejyense* Blechnum humbertii,
B. longepetiolatum, Blotiella coriacea, Deparia
marojejyensis (Tardieu) Kato, Elaphoglossum cer-
ussatum Tardieu, and E. marojejyense.

More recent studies of plant specimens collect-
ed in the Pare National (PN) de Marojejy prior to
the present expedition (cf. Humbert and collabo-
rators in 1959-1960; Guillaumet in 1972; Miller
and collaborators in 1988-1989) and deposited at
the herbarium of the Laboratoire de Phaneroga-
mic, MNHN, have added 61 taxa to Tardieu-

* Names of authors of the taxa mentioned in Table 2-
3 are not given in the text.

Blot's initial list (Rakotondrainibe, unpubl. data),
bringing the total number of pteridophytes to 168.
In contrast, 211 species and varieties of pterido-
phytes have been identified from the Reserve Spe-
ciale (RS) d'Anjanaharibe-Sud (Rakotondrainibe
& Raharimalala, 1998), which is about three times
smaller in area than the PN de Marojejy (18,255
vs. 60,050 ha) and is separated from it by a cor-
ridor of intact forest only about 35-40 km in
length (Fig. 2-1). This suggests that assessments
of pteridophyte species diversity at Marojejy were
substantially underestimated.

The present inventory of pteridophyte species
occurring along the eastern slope and the summit
area of the Marojejy Massif was conducted be-
tween 4 October and 20 November 1996. The
principal objective was to supplement the list of
pteridophytes known from the reserve and to
evaluate the density of the various taxa present
and their altitudinal distribution. An integrated
conservation and development program is cur-
rently being conducted in the Marojejy/ Anjana-
haribe-Sud reserves, and an assessment of the
most highly threatened species (whether because
of small population size or restricted geographic
distribution) is of potential value in this context.

Methodology

The present study was conducted using a
standardized methodology described earlier for
inventory work performed in 1994 at RS
d'Anjanaharibe-Sud (Rakotondrainibe & Rahari-

20

FIELDIANA: ZOOLOGY

malala, 1998) and PN d'Andohahela (Rakoton-
drainibe, 1999). Two phases were involved: (1)
an inventory of the pteridophytes present along
an altitudinal transect on the eastern slope running
between the lowest point in the reserve and its
summit, the entire length of which is exposed to
the trade winds, and (2) ecological surveys in a
series of 800 m 2 plots representative of the various
biotopes present, from which an evaluation can be
made of the diversity of the pteridophyte com-
munities present along the transect.

Floristic Survey

Five base camps were established along the
trail situated on the east slope and leading to the
summit area of the Marojejy Massif, between 400
m and 2132 m altitude: camp 1 at 450 m, camp
2 at 775 m, camp 3 at 1225 m, camp 4 at 1625
m, and camp 5 at 1875 m (see Chapter 1). The
inventory work was conducted within a radius of
2 km around each camp and along the main trail
from the point where it enters the park along the
banks of the Manantenina River, 3 km northwest
of the village of Mandena, to the summit (see
Chapter 1).

All voucher specimens are deposited at the her-
barium of the Departement des Recherches Fo-
restieres et Piscicoles, Centre National de la Re-
cherche Appliquee au Developpement DRFP/
FOFIFA) in Antananarivo (TEF), and the Labor-
atoire de Phanerogamie, MNHN. Duplicates of
most of the specimens collected will also be sent
to the Missouri Botanical Garden (MO) and the
Royal Botanic Gardens, Kew (K).

In most cases, generic delimitations follow those
proposed by Kramer and Green (1990), although for
certain genera whose circumscription is controver-
sial and which are in need of revision, particularly
for the Malagasy flora, more classical concepts have
been used: Thelypteridaceae follow the system of
Holttum (1974), and Grammitidaceae are classified
according to Pichi-Sermolli (1977), but the genus
Cochlidium has been separated from Xiphopteris ac-
cording to Bishop (1978). The identification of spe-
cies and varieties was done using monographs for
the genus Lastreopsis (Tindale, 1965), and the fam-
ilies Lindsaeaceae (Kramer, 1972) and Thelypteri-
daceae (Holttum, 1974). Circumscription of the oth-
er species follows treatments in the Flore de Mad-
agascar et des Comores (Stefanovic et al., 1997;
Tardieu-Blot, 1951-1971), except for Pyrrosia rho-
desiana, Ctenopteris flabelliformis, Elaphoglossum

ANTSIRAN'ANA

v\

oaA

/Efe'^m

-15S

f \ _C J Antalaha

/ 1 l s~t-*M T
\ Antananarivo / j

v w

/ V /

-20

^^^ <(/

I i

1 -''-N /

[ i

"fi

J/lfty

CfJ^yi

'~~/~

UiJ>

'-^. 1

^ \

^LJ*Sm

S y--J +* \

3

! /

25

Y p) Tolanaro

45

1

100 200km

50E

I

Fig. 2-1. Location of the PN de Marojejy (1), RS
d'Anjanaharibe-Sud (2), and PN d'Andohahela (3) in
Madagascar.

petiolatum ssp. salicifolium, and Lycopodiella car-
oliniana, which follow the most recent African flo-
ras (Schelpe, 1970; Schelpe & Diniz, 1979).

Ecological Sampling

Previous studies in Madagascar, at sites similar
to those found on the eastern slope of Marojejy
with intact or only slightly disturbed forest and
relatively uniform soil (Table 2-1), have shown
that altitude and topography influence the floristic
composition of pteridophyte communities (Rako-
tondrainibe, 1999; Rakotondrainibe & Guillerm,
1990; Rakotondrainibe & Raharimalala, 1988).

RAKOTONDRAINIBE: PTERIDOPHYTES

21

Table 2- 1 . Characteristics of the study plots on the eastern slopes of the PN de Marojejy.

Plot

Parameter

evaluated

PI

P2

P3

P4

P5

P6

Area (m : )

800

800

800

700

800

800

Altitude (m)

520

510

530

470

780

740

Topographic

middle

lower

plateau

middle

ridge

alluvial

position

slope

slope

slope

terrace

Exposure

S-SW

S-SE

N-NW

SW

N-E

Slope ()

10-15

5-10

5-10

45-50

15

0-2

Canopy height (m)

25

18-20

18-20

15-18

15-18

18-25

Woody plant cover

65

45-50

60

40-45

45-50

35-45

(%)

Herbaceous plant

15-20

20-50

10-15

7-10

10-15

20

cover (%)

Litter thickness

1-2

1-2

0.2-0.5

0.5-5

4-6

3

(cm)
Humus thickness

0-0.5

7-9

15-50

(cm)

Soil characteristics

yellow-

yellow-

yellow-

yellow-

yellow-

yellow-

ochre

ochre

ochre

ochre

ochre

ochre

clay

clay-sand

clay-sand

clay-sand

clay-sand

clay with
stones

Stratified sampling procedures were therefore
used in the present study: 18 plots covering a total
area of 14,100 m 2 were delimited along the alti-
tudinal transect, situated on slopes (plots PI -PI 5)
and along streams (R1-R3). The altitudinal and
topographic distributions of the plots, as well as
other physical and biological features, are pre-
sented in Tables 2-1 and 2-2. The locations of the
plots were not selected at random; instead, each
plot was established in an area in which both the
vegetation and the physical characteristics of the
milieu were considered to be homogeneous. Each
plot had a rectangular or near rectangular form.
The surface area of each plot was 800 m 2 , except
for P4 (700 m 2 ) and P13, P14, and P15 (600 m 2
each), which were situated in narrow or frag-
mented biotopes. The abundance of each species
was, however, always calculated (by extrapolation
if necessary) for a standard area of 800 m 2 . In
each plot, two kinds of information were record-
ed. First, species abundance was coded on a non-
linear scale according to the following categories:
1, a single individual or colony; 2, 2-4 individuals
or colonies; 3, 5-9 individuals or colonies; 4, 10-
19 individuals or colonies; 5, 20-49 individuals
or colonies, and 6, more than 50 individuals or
colonies. Second, the growth form of each species
was noted, as follows: T/r, always terrestrial, or
terrestrial and more rarely epilithtic; E/r, always

epiphytic, or epiphytic and more rarely epilithic;
R, strictly epilithic; L, lianescent; and T/E/r, ter-
restrial, epiphytic, or more rarely epilithic. The
criteria used to assess homogeneity within a plot,
the method used to count individuals and colo-
nies, and the definitions of the growth forms used
here are given in Rakotondrainibe and Rahari-
malala (1998). Epiphytic species in the upper lay-
ers of the forest were identified using binoculars,
and as a consequence, at least for the smallest
species in the families Hymenophyllaceae and
Grammitidaceae, their density was almost certain-
ly underestimated above 12-15 m.

Data Analysis

The floristic data collected from the plots were
treated using two complementary types of multi-
variate analyses (SPAD, version 3): correspon-
dence analysis (CA) and hierarchial ascending
classification (HAC) (Benzecri & Benzecri, 1984;
Lebart et al., 1997). Using a data matrix in which
species were plotted against sample plots, appli-
cation of these methods results in a double clas-
sification of columns (plots) and rows (species) in
which plots with similar species composition (and
therefore presumably similar environments) are
grouped together. The relations among the various

22

FIELDIANA: ZOOLOGY

Table 2-1. Extended.

Plot

P7

P8

P9

P10

Pll

P12

P13

P14

P15

800

800

600

800

800

800

600

600

600

820

1300

1290

1290

1540

1550

1530

1920

1900

middle

slope

middle/
lower
slope

ridge

higher
slope

lower
slope

ridge

middle
slope

ridge

lower
slope

N

N-NE

N

E-SE

NE

S

E

N

20

15-20

0-5

20-30

15

5-15

10-15

15

5-10

18-20

18

8-12

18

12-15

2-8

10-12

1-3

6-8

50-60

30-35

40-50

40-45

20-35

30-35

40

25-40

35-40

25

25-30

30-40

10 +
moss

35

30-50
(+ moss)

7

100 (moss)

40-50
(moss)

4-7
6-7

4-5
1

9-10

10-15

40-60

1-2
50-120

10-20
10-20

?
?

10-20
80

1-2
40

yellow-
ochre
clay-sand

yellow-
ochre
clay with
stones

yellow-
ochre
clay-sand

yellow-
ochre
clay-sand

?

White
clay-sand

yellow-
ochre
clay-
sand

elements of the matrix are measured using the x 2
distance (Lebart et al., 1997).

HAC generates a tree or dendrogram compris-
ing a hierarchical and progressive grouping of
plot units and/or species units in sets of increasing
size. The criterion used for grouping units into a
class is the average of the weighted distances. CA
generates scatter diagrams of plot and/or species
points represented in the plane of pair-wise sets
of axes resulting from the analysis. Each axis ex-
plains a percentage of the variance among the
points. The dispersion of the clusters of points
along a given axis is here an expression of an
ecological gradient.

The data matrix used for the two analyses list-
ed, in rows, all 194 species recorded in the plot
studies, against columns representing each of the
18 plots sampled (P1-P15 and R1-R3). Each cell
of the matrix contained the corresponding abun-

dance code (1-6) as defined above. The two anal-
yses assess clusters of plots and/or species, which
can then be interpreted with regard to their ecol-
ogy. The relative contribution of a species to the
variance expressed by a given axis will be higher
when a species is both abundant and present in a
limited number of plots. Species present in a clus-
ter of plots are referred to as "characteristic"
when their contribution to the fraction of the var-
iance expressed by the axis by which the group
was identified is ^1.8%. "Exclusively character-
istic" species are those that belong to a single
cluster of plots, and "preferentially characteris-
tic" species belong to at least two clusters. The
threshold value of 1.8% was chosen a posteriori
because it allows for the elimination of species
that are too widely distributed, those that are in-
frequent, and those whose distribution is erratic.
It should be noted that the concept of "character-

Table 2-2. Characteristics of PN de Marojejy study plots in forest located along streams.

Plot

Area (m 2 )

Width

of
stream-
Altitude (m) bed (m)

Nature of
streambed

Flow

Rl 800(200X4) 490-510 1-3 large granitic rocks + sand

R2 800(200X4) 750-810 2-6 large granitic rocks + sand

R3 800(200X4) 1290-1310 2-4 large granitic rocks + sand

gentle

variable with sudden flows

gentle

RAKOTONDRAINIBE: PTERIDOPHYTES

23

Table 2-3. Pteridophytes of PN de Marojejy: Floristic composition, habit type, and altitudinal distribution.

No.

Taxa

400-
550 m

740-
840 m

1250- 1500-
1350 m 2132 m

Habit
type

9
10
11

12
13
14
15
16
17

18

19

20
21

22
23
24
25
26
27
28

29

30
31

32

33
34
35
36
37
38
39
40
41

42

43
44
45
46
47
48
49
50
51

*Amauropelta bergiana (Schltdl.) Holttum
*Amauropelta sp. 1 (FR 3349)
*Angiopteris madagascariensis de Vriese
Antrophyum boryanum (Willd.) Kaulf.
*Antrophyum malgassicum C. Chr.
*Arthropteris monocarpa (H. L. Cordem.) C. Chr.
*Arthropteris orientalis (J. F. Gmel) Posth. var.

subbiaurita (Hook.) C. Chr.
*Arthropteris palisotii (Desv.) Alston
*Asplenium affine Sw. var. indet. (FR 3430)
Asplenium afzellii Rosend.
*Asplenium bipartitum Bory ex Willd.
*Asplenium cancellatum Alston
Asplenium cuneatum Lam.
*Asplenium dregeanum Kunze

* Asplenium friesiorum C. Chr.

Asplenium herpetopteris Baker var. herpetopteris

* Asplenium herpetopteris Baker var. acutipinna-
tum (Bonap.) Tardieu

*Asplenium herpetopteris Baker var. villosum
(Bonap.) Tardieu

*Asplenium inaequilaterale Willd.

Asplenium marojejyense Tardieu

*Asplenium nidus L.

Asplenium normale D. Don

Asplenium obscurum Blume

Asplenium pellucidum Lam.

"Asplenium petiolulatum Mett.

Asplenium sandersonii Hook.

* Asplenium thunbergii Kunze

Asplenium unilaterale Lam. et A. aff. unilaterale
(FR 3394)

Asplenium variabile Hook. var. paucijugum (F. Bal-
lard) Alson

"Asplenium sp. nov. 1 ined. (FR 3357)

Athyrium scandicinum (Willd.) C. Presl var. scandi-
cinum

* Athyrium scandicinum (Willd.) C. Presl var. bipin-

nata Rakotondr.
"Belvisia spicata (L.f.) Mirb.
"Blechnum attenuatum (Sw.) Mett.
*Blechnum bakeri (Baker) C. Chr.
Blechnum humbertii Tardieu
"Blenchum ivohibense C. Chr.
Blechnum longepetiolatum Tardieu.
Blechnum madagascariense Tardieu.
*Blechnum simillimum (Baker) Diels
"Blechnum simillimum (Baker) Diels fa. binerve

(Hook.) Tardieu
"Blechnum simillimum (Baker) Diels var. xipho-

phyllum (Baker) Tardieu
Blotiella coriacea (Tardieu) Rakotondr. comb. nov.
*Blotiella madagascariensis (Hook.) Tryon
Bolbitis auriculata (Lam.) Alston
*Bolbitis humblotii (Baker) Ching
"Cheilanthes sp. nov. 1 ined. (FR 3621, 3692)
Christella dentata (Forssk.) Brownsey & Jermy
Christella distans (Hook.) Holttum
*Cochlidium serrulatum (Sw.) L. E. Bishop
Coniogramme madagascariensis C. Chr.

-

-

+

+

T

+

+

-

-

T

+

-

-

-

T

+

+

+

-

E

+

+

-

-

E

-

+

+

-

T/E/r

+

+

-

E

+

-

-

E

-

+

-

-

T

+

+

+

+

T/E

+

+

-

-

E

+

+

-

-

E

+

+

-

-

E/r

+

+

-

-

E/r

-

-

+

-

E

+

+

+

+

E/r

+

+

-

-

E

+

+

-

-

E

+

+

-

-

T/r

-

+

-

-

T/r

+

+

-

-

E

-

-

+

+

T

-

+

-

-

T/r

+

+

+

-

E/r

-

+

-

-

E

-

+

+

-

E

-

+

-

-

T/E

+

+

+

-

T/r

+

-

-

-

T/E

+

-

-

-

E/r

+

+

T

-

-

+

+

T

+

+

+

+

E

-

+

+

-

T/E/r

-

-

+

-

R

-

-

+

+

T

-

-

+

+

T

-

-

-

+

T

-

-

-

+

T

+

+

+

+

T/E/r

+

T/r

-

-

+

-

T

-

-

+

+

T

-

+

+

+

E

+

+

-

-

T/r

-

+

-

-

T

-

-

-

+

T

+

-

-

-

T

+

+

-

-

T

-

+

+

+

E

-

+

-

T

24

FIELDIANA: ZOOLOGY

Table 2-3. Continued.

No. Taxa

400-

740-

1250-

1500-

Habit

550 m

840 m

1350 m

2132 m

type

-

+

+

-

T

+

-

T

+

-

-

-

T

-

-

+

-

T

-

-

+

+

E

-

+

+

+

E

+

+

+

-

E

+
+

E
E

-

-

+

+

E

-

-

+

+

E

-

+

+

-

E

-

-

+

-

T

-

-

-

+

T

-

-

+

+

T

-

-

+

+

T

+

+

T

+

-

-

-

T

-

+

+

+

T

-

+

-

-

T

+

+

+

T

-

-

+

-

T

+

+

+

+

T

-

+

+

+

T

-

-

-

+

T

+

+

-

-

T

+

-

-

-

T

+

+

+

-

T

-

-

+

-

T

-

+

-

-

T

+

T

+

+

-

-

E

+

+

-

-

E

+

+

+

-

T

-

+

-

-

T

-

+

+

+

T

-

+

-

-

T

-

-

+

-

T

-

+

-

-

T

+

+

-

-

T

+

+

-

-

T

+

-

-

-

E

-

+

+

-

T

-

-

-

+

E

-

-

+

+

T/E

+

T

-

-

+

+

T/E

-

+

-

-

E

-

-

+

+

E

52 *Ctenitis ochrorachis (Baker) Tardieu var. violacea

Rakotondr.

53 *Ctenitis poolii (C. Chr.) Tardieu

54 *Ctenitis warburii (C. Chr.) Tardieu

55 *Ctenitis sp. nov. 1 ined. (FR 3558)

56 Ctenopteris alboglandulosa (Bonap.) Tardieu

57 *Ctenopteris devoluta (Baker) Tardieu

58 ^Ctenopteris elastica (Bory) Copel.

59 * Ctenopteris flabelliformis (Poir.) J. Sm.

60 Ctenopteris humbertii (C. Chr.) Tardieu

61 ^Ctenopteris sp. 1 cf. C. humbertii (FR 3484, 3615

bis)

62 * Ctenopteris villosissima (Hook.) Harley

63 *Ctenopteris zenkeri (Hieron.) Tardieu

64 *Cyathea aff. albida Tardieu

65 Cyathea alticola (Tardieu) Tindale

66 *Cyathea approximata Bonap.

67 *Cyathea bellisquamata Bonap. var. basilobata C.

Chr.

68 *Cyathea boivinii Mett.

69 *Cyathea borbonica Desv. var. ? (FR 3282)

70 * Cyathea bullata (Baker) Rakotondr.

71 *Cyathea bullata (Baker) Rakotondr. var. mada-

gascarica (Bonap.) Rakotondr.

72 *Cyathea costularis Bonap.

73 *Cyathea coursii (Tardieu) Rakotondr.

74 * Cyathea decrescens Mett.

75 * Cyathea dregei Kunze

76 *Cyathea dregei Kunze var. polyphlebia C. Chr.

77 *Cyathea ligulata Tardieu

78 *Cyathea longipinnata Bonap.

79 *Cyathea melleri (Baker) Domin var. melleri

80 *Cyathea perrieriana C. Chr.

8 1 * Cyathea serratifolia Baker et C. aff. serratifolia

(FR 3379)

82 *Cyclosorus interruptus (Willd.) H. Ito

83 *Davallia chaerophylloides (Poir.) Steud.

84 Davallia repens (L.f.) Kuhn (= Humata repens

(L.f.) Diels)

85 *Deparia parvisora (C. Chr.) M. Kato et D. aff.

parvisora (FR 3346)

86 *Deparia sp. 1 (FR 3445, 3447) et var. (FR 3446)

87 *Dicranopteris linearis (Burm.f.) Underw.

88 Didymochlaena microphylla (Bonap.) C. Chr.

89 *Diplazium andapense (Tardieu) Rakotondr.

90 *Diplazium dilatatum Blume

91 Diplazium nemorale (Baker) Schelpe

92 * Diplazium proliferum (Lam.) Thouars

93 *Drynaria willdenowii (Bory) T Moore

94 *Dryopteris mangindranensis Tardieu

95 * Elaphoglossum achroalepis (Baker) C. Chr.

96 Elaphoglossum acrostichoides (Hook. & Grev.)

Schelpe

97 * Elaphoglossum angulatum (Blume) T Moore

98 * Elaphoglossum aubertii (Desv.) T Moore

99 * Elaphoglossum decaryanum Tardieu

100 * Elaphoglossum deckenii (Kuhn) C. Chr. var. rufi-

dulum (Willd. ex Kuhn) Tardieu

101 Elaphoglossum forsythii-majoris H. Christ

RAKOTONDRAINIBE: PTERIDOPHYTES 25

Table 2-3. Continued.

No.

Taxa

400-
550 m

740- 1250- 1500-
840 m 1350 m 2132 m

Habit
type

7 (FR 3574, 3582. 3607)
10 (FR 3485, 3488. 3489,

102 * Elaphoglossum humbertii C. Chr.

103 Elaphoglossum lepervanchii (Fee) Moore

104 * Elaphoglossum leucolepis (Baker) Krajina ex

Tardieu

105 Elaphoglossum marojejyense Tardieu

106 * Elaphoglossum petiolatum (Sw.) Urb. ssp. salici-

folium (Willd. ex Kaulf.) Schelpe

107 * Elaphoglossum scolopendriforme Tardieu et E.

aff. scolopendriforme

108 Elaphoglossum aff. sieberi (Hook. & Grev.) T.

Moore

109 Elaphoglossum spathulatum (Bory) T Moore

1 10 * Elaphoglossum subsessile (Baker) C. Chr.

1 1 1 * Elaphoglossum sp. 5 (FR 3458. 3472. 3475.

3611. 3701)

1 12 * Elaphoglossum sp. 6 (FR 3496. 3514. 3520,

3550. 3616)

1 13 * Elaphoglossum sp,

1 14 * Elaphoglossum sp,

3646. 3408)

1 15 * Elaphoglossum sp. 11 (FR 3468. 3593)

1 16 *Gleichenia madagascariensis C. Chr.

1 17 *Gleichenia polypodioides (L.) J. E. Smith.

1 18 Grammitis barbatula (Baker) Copel.

1 19 *Grammitis copelandii Tardieu

120 Grammitis cryptophlebia (Baker) Copel.

121 *Grammitis ebenina (Maxon) Tardieu

122 Grammmitis gilpinae (Baker) Tardieu

123 Grammitis holophlebia (Baker) Copel.

124 *Grammitis kyimbilensis (Brause) Copel.

125 *Grammitis microglossa (C. Chr.) Criing

126 *Grammitis obtusa Willd. ex Kaulf.

1 27 Grammitis sxnsora (Baker) Copel.

128 *Grammitis sp. 1 (FR 3589. 3629. 3668. 3668 bis)

129 *Grammitis sp. 3 (FR 3663)

130 *Histiopteris incisa (Thunb.) J. Sm.

131 *Huperzia cavifolia (C. Chr.) Tardieu

132 *Huperzia megastachya (Baker) Tardieu

133 *Huperzia obtusifolia (P. Beauv.) Rothm.

134 *Huperzia ophioglossoides (Lam.) Rothm.

135 *Huperzia pecten (Baker) Tardieu

136 *Huperzia suberecta (Lowe) Tardieu

137 *Huperzia squarrosa (G. Forst.) Trevis.

138 *Huperzia trigona (C. Chr.) Tardieu

139 *Huperzia verticillata (L.f.) Trevis.

140 *Hymenophyllum capillare Desv. var. ? (FR 3487)

141 * Hymenophyllum deltoid cum C. Chr.

142 *Hymenophyllum fumarioides Willd. et H. aff. fu-

marioides (FR 3365. 3652. 3700)

143 *Hymenophyllum heimii Tardieu

144 Hymenophyllum hirsutum (L.) Sw.

145 * Hymenophyllum humbertii C. Chr.

146 * Hymenophyllum inaequale (Poir.) Desv.

147 * Hymenophyllum parvum C. Chr. (= H. capense

Schrad.)

148 * Hymenophyllum polyanthos (Sw.) Sw.

149 * Hymenophyllum poolii Baker

150 * Hymenophyllum sibthorpioides Mett.

151 * Hymenophyllum tunbrigense (L.) Smith

-

-

+

+

E

-

+

-

+

T/E

+

+

+

E

-

+

-

-

E

+

-

-

E/r

-

+

+

+

E

+

+

+

+

E

+

-

-

-

R

-

+

+

+

T/E

-

+

+

+

E

Ti-;

-

-

+

+

E

-

+

+

+

E

-

-

+

+

T/E

-

-

-

+

T

-

-

-

+

T

-

-

+

-

E

-

-

+

+

E

-

-

-

+

E

-

-

+

+

E

-

+

+

+

E

-

-

+

+

E

-

+

+

-

E

-

-

-

+

E

-

-

-

+

E

-

+

-

-

E

-

-

+

+

E

-

-

-

+

E

-

-

-

+

T

+

+

+

-

E

+

+

+

+

T/E

-

-

-

+

T

+

+

-

+

E

-

+

+

-

E

-

-

-

+

R

-

-

+

+

E

-

-

+

+

E

-

-

+

+

E

-

-

+

+

E

-

-

+

+

E

-

-

+

T/E

-

-

+

+

E

-

+

+

+

T/E/r

-

-

+

+

T/E

-

-

+

+

E

+

+

+

E

-

+

+

+

E

-

-

+

+

T/E

-

+

+

+

T/E

-

+

+

E

26

FIELDIANA: ZOOLOGY

Table 2-3. Continued.

No.

Taxa

400-
550 m

740-
840 m

1250- 1500-
1350 m 2132 m

Habit
type

152 *Hymenophyllum veronicoides C. Chr.

153 *Hymenophyllum viguieri Tardieu

154 *Lastreopsis subsimilis (Hook.) Tindale

155 Lepisorus excavatus (Bory ex Willd.) Moore

156 *Lindsaea blotiana K. U. Kramer

157 *Lindsaea coursii (Tardieu) K. U. Kramer

158 *Lindsaea ensifolia Sw.

159 Lindsaea flabellifolia (Baker) Kuhn

160 Lindsaea goudotiana (Kunze) Kuhn

161 * Lindsaea madagascariensis Baker

162 * Lindsaea millefolium K. U. Kramer

163 * Lindsaea odontolabia (Baker) K. U. Kramer

164 Lindsaea odorata Roxb.

165 * Lindsaea sp. nov. 1 (FR 3605)

166 *Lomariopsis crassifolia Holttum

167 Loxogramme humblotii C. Chr.

168 Loxogramme lanceolata (Sw.) C. Presl. et L. aff.

lanceolata (FR 3645)

169 *Lycopodiella caroliniana (L.) Pic. Serm.

170 *Lycopodiella cernua (L.) Pic. Serm.

171 *Lycopodium zanclophyllum Wilce

172 *Lygodium lanceolatum Desv.

173 *Macrothelypteris torresiana (Gaudich.) Ching

174 *Marattia fraxinea Sm. ex J. F. Gmel.

175 Microlepia madagascariensis C. Presl

176 Microsorum punctatum (L) Copel.

177 *Monogramma graminea (Poir.) Schkuhr

178 Nephrolepis biserrata (Sw.) Schott

179 *Nephrolepis tuberosa (Bory) C. Presl

180 Odontosoria melleri (Hook.) C. Chr.

181 Oleandra distenta Kunze

1 82 Ophioglossum palmatum L.

183 Ophioglossum pendulum L.

184 *Pellaea angulosa (Bory) Baker

185 Phymatosorus scolopendria (Burm. f.) Pic. Serm.

186 *Pityrogramma calomelanos (L.) Link

1 87 *Pityrogramma humbertii C. Chr.

188 Pityrogramma humbertii C. Chr. var. ? (FR 3687)

189 *Platycerium madagascariense Baker

190 *Pleopeltis macrocarpa (Bory ex Willd.) Kaulf.

191 Pneumatopteris remotipinna (Bonap.) Holttum

192 Pneumatopteris subpennigera (C. Chr.) Holttum

193 Pseudocyclosorus pulcher (Bory ex Willd.) Holt-

tum

194 *Pseudophegopteris cruciata (Willd.) Holttum

195 *Pteris catoptera Kunze et P. aff. catoptera (FR

3584)

196 *Pteris cretica L.

197 *Pteris elongatiloba Bonap. var. remotivenia Bon-

ap.

198 *Pteris pseudolonchitis Bory ex Willd.

199 *Pteris woodwardioides Bory ex Willd.

200 *Pyrrosia rhodesiana (C. Chr.) Schelpe

201 Rumohra adiantiformis (G. Forst.) Ching

202 Rumohra lokohoensis Tardieu

203 Saccoloma henriettae (Baker) C. Chr.

204 *Schizaea dichotoma L.

205 *Schizaea pectinata (L.) Sw.

206 *Selaginella fissidentoides (Hook. & Grev.) Spring

fa fissidentoides

-

-

+

-

E

+

-

-

+

E

-

-

+

-

T

-

+

+

+

E

-

+

+

+

T

-

-

+

+

E

-

+

-

-

T

-

+

+

-

T

-

-

+

-

E

-

-

-

+

E

-

+

-

-

T/E

-

+

-

-

E

-

-

-

+

R

-

-

-

+

T/E

-

+

-

-

T/E

+

+

-

-

E/r

+

+

+

+

E

-

-

-

+

T

-

+

+

+

T

-

-

-

+

T

+

-

-

-

L

+

+

-

-

T

+

+

+

-

T

+

+

+

-

T

+

+

+

-

E

+

+

-

-

E

+

+

-

-

T/E/r

-

+

+

-

E

-

+

-

+

T

+

+

+

-

E

-

-

+

-

E

+

-

-

-

E

-

+

-

-

T

+

+

+

-

E

+

-

-

-

T

-

-

-

+

T

-

-

-

+

T

-

+

+

-

E

+

-

-

+

E

+

+

+

+

T

-

+

+

-

T

-

+

-

-

T

-

+

-

-

T

+

+

+

-

T

-

+

T

+

-

-

-

T

-

+

_

T

+

-

-

-

T

-

+

-

-

E

+

+

+

+

E

-

-

+

+

T/E

-

-

+

+

T

+

+

-

-

T

-

-

-

+

T

+

+

-

-

T

RAKOTONDRAINIBE: PTERIDOPHYTES

27

Table 2-3. Continued.

No.

Taxa

400-
550 m

740-
840 m

1250- 1500-
1350 m 2132 m

Habit
type

207 *Selaginella goudotana Spring van goudotana

208 *Selaginella hildebrandtii A. Braun ex Hieron.

209 *Selaginella pervillei Spring

210 *Selaginella polymorpha Badre (= S. pectinata

Spring)

2 1 1 *Selaginella unilateralis Spring

212 Sphaerostephanos arbuscula (Willd.) Holttum

213 *Sphenomeris chinensis (L.) Maxon

214 *Stenochlaena tenuifolia (Desv.) Moore

215 *Sticherus flagellaris (Bory) St John

216 Tectaria gemmifera (Fee) Alston et T. aff. gemmi-

fera (FR 3281)

2 1 7 Tectaria humbertiana Tardieu

218 *Tectaria magnifica (Bonap.) Tardieu

219 *Trichomanes bipunctatum Poir.

220 *Trichomanes bonapartei C. Chr.

221 *Trichomanes borbonicum Bosch

222 Trichomanes cupressoides Desv.

223 *Trichomanes digitatum Sw.

224 *Trichomanes erosum Willd.

225 *Trichomanes lenormandii Bosch

226 *Trichomanes longilabiatum Bonap.

227 *Trichomanes meifolium Bory ex Willd.

228 *Trichomanes montanum Hook.

229 ^Trichomanes montanum Hook. var. ? (FR 3339)

230 ^Trichomanes rigidum Sw.

231 Trichomanes speciosum Willd.

232 Vittaria ensiformis Sw.

233 *Vittaria humblotii Hieron.

234 Vittaria scolopendrina (Bory) Thwaites

235 Xiphopteris mysuroides (Sw.) Kaulf.

236 *Xiphopteris oosora (Baker) Alston var. micropec-

ten C. Chr.

237 *Xiphopteris sikkimensis (Hieron.) Copel.

238 *Xiphopteris sp. 3 (FR 3613, 3680)

239 * Xiphopteris sp. 4 (FR 3519)

-

+

-

-

T

-

+

-

-

T/r

+

-

-

-

T

+

+

+

T

+

+

T

-

+

-

-

T

-

+

-

-

T

+

+

-

-

T/E

-

+

+

+

T

+

-

-

-

T

+

-

-

T

-

+

-

-

T

+

+

+

-

E

+

+

+

+

E

-

+

+

+

E/r

+

+

+

+

T/r

-

+

+

+

E

+

+

+

+

E

+

+

+

-

E

+

-

-

-

R

-

+

+

+

T/E

-

+

+

-

E

+

-

+

-

E/r

-

+

+

+

T

+

-

+

-

L

+

+

-

-

E

-

+

+

-

E

+

-

-

-

E

-

-

-

+

E

-

-

-

+

E

_

+

E

-

-

-

+

E

-

-

+

-

E

Species endemic to Madagascar are indicated in boldface type. Species endemic to PN de Marojejy are indicated
in boldface type and underlined .

* New records for PN de Marojejy.

T = terrestrial; E = epiphytic; R = strictly epilithic; r = occasionally epilithic; L = lianescent; + = present; - =
absent; FR = initials of the collector's name, France Rakotondrainibe.

istic" species in the present context is based on
the ecological amplitude of one species and its
abundance, rather than on its constancy in a veg-
etation unit, as often utilized by phytosociologists.

Results

General Floristic Inventory

Species Richness Table 2-3 lists all the taxa
observed on the eastern slope and in the summit

area of the PN de Marojejy, inside and outside the
18 plots, between 400 and 2132 m altitude, and
indicates the growth form (terrestrial, epiphytic,
or epilithic) and altitudinal distribution of each
taxon. The delimitation of the altitudinal zones is
explained below.

A total of 239 species and varieties of pteri-
dophytes, representing 70 genera, were identified.
The most speciose genera are Asplenium (20 spp.
and 2 varieties), Elaphoglossum (21 spp.), Cy-
athea (16 spp. and 2 varieties), Hymenophyllum
(14 spp.), Trichomanes (13 spp. and 1 variety),
Grammitis (12 spp.), Lindsaea (10 spp.), Blech-

28

FTELDIANA: ZOOLOGY

Fig. 2-2. Percentages of the whole pteridophyte flora
(239 species and varieties listed in Table 2-3) by habit
type in the PN de Marojejy. T/r = always terrestrial or
terrestrial and more rarely epilithic; E/r = always epi-
phytic or epiphytic and more rarely epilithic; R = strict-
ly epilithic; L = lianescent; T/E/r = terrestrial, epiphytic
or more rarely epilithic.

num (7 spp. and 2 varieties), Huperzia (9 spp.),
and Ctenopteris (8 spp.). The 171 taxa marked
with an asterisk in Table 2-3 are new records for
the PN de Marojejy.

Endemism The 105 reserve species endemic
to Madagascar are indicated in Table 2-3 by bold
characters; they represent 43.9% of the pterido-
phyte flora of the park. An additional 18 species
are endemic to the Malagasy region, which in-
cludes Madagascar, the Mascarenes (Reunion,
Mauritius, and Rodrigues), the Seychelles, and the
Comoro Islands, bringing the total number of re-
gional endemics to 123 taxa (51.5% of the total).

Among the species endemic to the Marojejy
Massif, only six have been observed during the
present expedition on the eastern slope: Blechnum
longepetiolatum, Cheilanthes sp. nov. 1, Cyathea
alticola, Elaphoglossum sp. nov. 1 1 , Lindsaea sp.
nov. 1, and Tectaria humbertiana. Several other
endemic species were collected by Humbert on
the western slope of the massif and nearby areas,
including Asplenium andapense Tardieu, Deparia
marojejyensis (Tardieu) Kato, Diplazium maroje-
jy ense (Tardieu) Rakotondr., and Elaphoglossum
cerussatum Tardieu. Among the other species
considered by Tardieu as endemic to Marojejy,
Blotiella coriacea was recorded by Perrier de la
Bathie (Bathie, no. 15297P!) in the Tsaratanana
Massif, and Asplenium marojejyense, Blechnum
humbertii, and Elaphoglossum marojejyense have
since been recorded by the present author in other
reserves within the country (Rakotondrainibe &
Quansah, 1994; Rakotondrainibe & Raharimalala,

c

t

D

C
B

P12

P13

P9

P11

P15

P14

R3 =

P8

P1Q_

P5

P7

P6

R2

R1

P2

P3

P4

P1

Fig. 2-3. Hierarchical ascending classification
(HAC) tree using a matrix of 18 plots (P1-P15, R1-R3)
and 194 species of pteridophytes recorded in the PN de
Marojejy. The criterion used for grouping units into a
class is the average of the weighed distances.

1996, 1998). One widespread species, Arthropter-
is palisotii, is currently known in Madagascar
only from the Marojejy Massif, but additional
populations occur in West Africa, the Comoros,
Sri Lanka, Java, and several areas in Oceania
(Tardieu-Blot, 1964).

Growth Forms The percentage of each habit
type represented in Figure 2-2 was calculated us-
ing the data in Table 2-3. Approximately equal
percentages of pteridophyte species and varieties
are either (1) terrestrial or more rarely epilithic
(T/r) (42.7%) or (2) epiphytic or epiphytic and
more rarely epilithic (E/r) (43.9%); 10.5% of the
taxa are variously terrestrial, epiphytic, or epilith-
ic (T/E/r), whereas strictly epilithic (R) (2.1%) or
lianescent species (L) (0.8%) are rare.

Distribution of Taxa

Ecological Gradients and Characteristic
Taxa The tree obtained by HAC from the data
matrix (not given here) of the study plots (Pl-
P15 and R1-R3) against the 194 species recorded
collectively in all the plots is presented in Figure
2-3. By sectioning the tree between nodes 32 and
33, four groups of plots are defined on the basis
of their floristic composition: group A (PI, P2, P3,
P4, Rl), group B (P5, P6, P7, R2), group C (P8,
P10, R3), and group D (P9, Pll, P12, P13, P14,
P15).

Using the positions of groups A through D,

RAKOTONDRAINIBE: PTERIDOPHYTES

29

II [Xj = ll.

64%)

Fig. 2-4. Correspondence analysis (CA) of the matrix of 18 plots and 194 pteridophyte species recorded in the
PN de Marojejy: projection on axes I and II of all the plots (PI -PI 5, R1-R3). Group A = plots at low altitude (470-
530 m); group B = plots at middle altitude (740-820 m); group C = plots at middle altitude (1290-1310 m); group
D = plots at high altitude (1530-1920 m).

axes I II (Fig. 2-4), I III (Fig. 2-5), and I-IV
(Fig. 2-6), and the ecological information given
in Table 2- 1 and 2-2, it is possible to evaluate the
ecological gradients responsible for the floristic
heterogeneity among pteridophytes on the eastern
slope of the Marojejy Massif. Axes I, II, and IV
show the same altitudinal gradient, whereas axis
III reflects a topographic gradient that is striking
but localized. Along axis I, group A, comprising
plots at low altitude (470-530 m), is separated

from group D, made up of plots at high altitude
(1530-1920 m), whereas groups B (740-820 m)
and C (1290-1310 m), at middle altitudes, occupy
an intermediate position on this axis. Axis II sep-
arates groups A and B and groups C and D, re-
spectively. The projection of plot points on axes
I-IV shows fine-scale heterogeneity among the
plots constituting group D, which can be further
divided into two subgroups, Dl (P9, PI 2, PI 3)
and D2 (PI 1, P14, P15), as indicated in Figure

III (^=9.22%)

19.10%)

Fig. 2-5. Correspondence analysis (CA) of the matrix of 18 plots and 194 pteridophytes species recorded in the
PN de Marojejy: projection on axes I and III of all the plots (P1-P15, R1-R3). Group A = plots at low altitude
(470-530 m); group B = plots at middle altitude (740-820 m); group C = plots at middle altitude (1290-1310 m);
group D = plots at high altitude (1530-1920 m).

30

FIELDIANA: ZOOLOGY

IV (}.4=8.05%)

Fig. 2-6. Correspondence analysis (CA) of the matrix of 18 plots and 194 pteridophytes species recorded in the
PN de Marojejy: projection on axes I and IV of all the plots (P1-P15, R1-R3). Group A = plots at low altitude
(470-530 m); group B = plots at middle altitude (740-820 m); group C = plots at middle altitude (1290-1310 m);
group D = plots at high altitude (1530-1920 m).

2-6 and in the dendrogram of Figure 2-3. In Fig-
ure 2-5 the parcels belonging to group B are scat-
tered along axis III in relation to their topographic
position: plots P7 and P5, situated respectively on
a ridge and mid-slope, have negative values; plot
R2, located along the banks of a creek, has a pos-
itive value; plot P6, however, on a raised alluvial
terrace, has an intermediate value. It should be
noted that the position of plot P9 on both the tree
and the two-dimensional projections is not con-
sistent with the overall pattern of altitudinal dis-
tribution described above. Instead, although this
plot is located at 1290 m, it clearly belongs to
group D on the basis of its floristic composition.
The distinctive topographic position of plot P9, on
a ridge that is exposed to the elements, is probably
sufficient to explain this anomaly. This plot was
established on a very narrow, straight, nearly flat
ridge with abrupt slopes on both sides that further
accentuate the ridge effect. Moreover, the physi-
ognomy of the vegetation in P9 is exceptional for
this altitude; it is a low forest about 8-12 m tall
(Table 2-1) and comprised of trees whose twisted
branches are covered with epiphytes.

On the basis of floristic composition of the pte-
ridophytes in the plots, four altitudinal zones can
thus be recognized on the eastern slope of the PN
de Marojejy (as indicated in the column headings
of Table 2-3):

Zone A, at low altitude, studied at plots between
400 and 550 m (PI, P2, P3, P4, Rl)

Zone B, at lower mid-altitude, studied between

740 and 840 m (P5, P6, P7, R2)
Zone C, at upper mid-altitude, studied between

1250 and 1350 m (P8, P10, R3)
Zone D, at high altitude, studied between 1500

and 1900 m (P9, Pll, P12, P13, P14,

P15)

The sampling method used did not, however, per-
mit a precise determination of the upper and lower
altitudinal limits of these zones. On the other
hand, it was possible to detect the influence of
topography on the species composition of the pte-
ridophytes, which appeared to be restricted to
zone B, between 740 and 840 m.

Plot P9, at 1290 m on a ridge, has been includ-
ed in zone B on the basis of its floristic compo-
sition, even though it occurs considerably higher
on Marojejy. This reassignment only emphasizes
how difficult it is to establish fixed altitudinal lim-
its between floristic zones whose distribution is
influenced not only by altitude but also by local
topographic conditions.

The 49 characteristic species (including both
exclusively and preferentially characteristic spe-
cies) for each of the altitudinal and/or topographic
zones (ridges, mid-slopes, bottomlands, and creek
banks) are listed in Table 2-7. As indicated above,
each of these species contributes at least 1.8% to
the variance of one or more of the four axes de-
fined in the factorial analysis, in particular to axes

RAKOTONDRAINIBE: PTERIDOPHYTES

31

Table 2-4. Percentages of the pteridophyte flora
(239 species and varieties listed in Table 2-3) by habit
types for each altitudinal zone in PN de Marojejy.

Table 2-6. Species richness for each altitudinal
zone of the 10 largest pteridophyte genera in PN de
Marojejy.

Zone A Zone B Zone C

Zone D

Zone

Zone

Zone

Zone

Habit 400- 740- 1250-

1500-

A

B

C

D

Total

type 550 m 840 m 1350 m

2132 m

400-

740-

1250-

1500-

400-

550

840

1350

2132

2132

T/r 40.2 43.1 32.8
E/r 47.5 45.4 52.5

34.5
48.7

Genus

m

m

m

m

m

T/E/r 7.3 11.5 13.1

15.0

Asplenium

14

18

7

3

22

R 2.5 0.0 0.8

1.0

Elaphoglossum

2

10

13

17

21

L 2.5 0.0 0.8

0.0

Cyathea
Hymenophyllum

7
1

9

4

9

12

8
13

18

14

T/r = always terrestrial or terrestrial and i

nore rarely

Grammitis

3

8

10

12

epilithic; E/r = always epiphytic or epiphytic and more

Trichomanes

8

10

12

7

13

rarely epilithic; R = strictly epilithic; L = lianescent; T/

Lindsaea

5

4

5

10

E/r = terrestrial, epiphytic, or more rarely epilithic.

Huperzia

3

4

6

7

9

Blechnum

1

2

6

6

9

Ctenopteris

1

3

6

6

8

I, II, and IV for the altitudinal indicator species
and to axis III for the topographic indicators.

Floristic Characteristics of the Altitudinal
Zones Percentages of the pteridophyte flora
(239 species and varieties listed in Table 2-3) by
habit type for each altitudinal zone are indicated
in Table 2-4. Table 2-5 shows the total number of
species and genera of the whole pteridophyte flora
(i.e., the generic and overall species richness) for
each altitudinal zone, along with the number and
percentage of species endemic to Madagascar and
the mean number of species per 800 m 2 plots (spe-
cies density). Table 2-6 lists the largest genera and
the number of species and varieties belonging to
them that occur within each altitudinal zone. The
highest generic and species richness occurred in
zone B (56 genera and 130 species), whereas the
greatest species density was observed at a some-
what higher altitude, in zone C (60 species/800
m 2 ). The level of species endemism increases pro-
gressively with altitude, from 36.6% in zone A to
a maximum of 49.5% in zone D. The percentage
of epiphytic species is always higher than for ter-
restrial taxa, regardless of altitude, and reaches a
maximum value between 1250 and 1350 m in
zone C. However, the percentage of species that

can be either epiphytic or terrestrial increases with
altitude, as the moss layer on the ground becomes
more widespread and thicker. Above 1500 m, and
especially near 1800 m, the moss coat present on
tree trunks and branches is continuous with the
thick layer on the ground, sometimes making it
difficult to distinguish between epiphytic and ter-
restrial species. Above 2000 m the forest gives
way to a high-altitude xerophytic grassland in
which all the fern species are terrestrial.

The generic and species composition of the pte-
ridophyte flora on the eastern slope of Marojejy
changes with altitude, as follows (cf. Tables 2-6
and 2-7):

Between 400 and 550 m, the genera Asplenium
(14 spp.), Trichomanes (8 spp.), and Cyathea
(7 spp.) are the most speciose. Three exclusive-
ly characteristic species occur, as well as three
preferentially characteristic species.

Between 740 and 840 m, the same three genera
are represented by a large number of species,
Asplenium (18 spp.), Trichomanes (10 spp.),
and Cyathea (9 spp.), along with Elaphoglos-

Table 2-5. Generic richness, species richness, and endemism rate of the pteridophyte flora (239 species listed in
Table 2-3) and mean number of species per 800 m 2 plots, for each altitudinal zone in PN de Marojejy.

Character

Zone A
400-550 m

Zone B
740-840 m

Zone C Zone D

1250-1350 m 1500-2132 m

Number of genera

Total number of species

Number of Malagasy-endemic species

Percentage of Malagasy-endemic species

Mean number of species/800 m 2

41

56

43

33

82

130

122

113

30

49

55

55

36.6

37.7

45.1

49.5

30.2

48.0

60.0

38.3

32

FIELDIANA: ZOOLOGY

sum (10 spp.)- There are many more exclusively
characteristic species (9) and preferentially
characteristic species (8).

Between 1250 and 1350 m, Elaphoglossum is
even more speciose (13 spp.), as are Hymeno-
phyllum (12 spp.) and Trichomanes (12 spp.);
the genera Cyathea (9 spp.) and Grammitis (8
spp.) are also well represented. Only one exclu-
sively characteristic species is present, but 6
preferentially characteristic species were re-
corded.

Between 1500 and 2132 m altitude, Elaphoglos-
sum (17 spp.) is particularly diversified, along
with Hymenophyllum (13 spp.) and Grammitis
(10 spp.). Five exclusively characteristic spe-
cies and 2 preferentially characteristic species
are found here.

Zone C, between 1250 and 1350 m, exhibits
features typical of a transition zone, with a greater
number of preferentially characteristic species
than exclusively characteristic species. Zone C
also shows a sharp decrease in the number of taxa
belonging to Asplenium, which is a dominant ge-
nus at lower altitudes, coupled with a marked in-
crease in species diversity in Elaphoglossum and
Hymenophyllum, both of which are especially
well represented at higher altitudes in zone D.

Discussion

Sampling Quality Rare Taxa

A total of 239 species and varieties of pteri-
dophytes were recorded on Marojejy. Of that to-
tal, 194 taxa (81.2%) were present in one or more
of the 18 sample plots examined. Among the 45
remaining pteridophytes, observed only outside
the plots, some are rare within the reserve or even
in Madagascar as a whole, including Asplenium
ajfine, A. obscurum, Bolbitis humblotii, Cyathea
borbonica var. ?, C. bullata var. madagascarica,
C. aff. albida, C. longipinnata, C. serratifolia,
Drynaria wildenowii, Elaphoglossum forsythii-
majoris, E. spathulatum, Grammitis barbatula,
Huperzia obtusifolia, and Selaginella goudotana
each of which was seen only once or twice. Sev-
eral other rare pteridophytes are known from only
one to four localities in the entire country, includ-
ing Blechnum simillimum var. xiphophyllum,
Cheilanthes sp. nov. 1, Ctenitis warburii, C. sp.
nov. 1, Ctenopteris sp. aff. humbertii, Diplazium

dilatatum, Grammitis microglossa, Huperzia sau-
rurus, Lastreopsis subsimilis, Lindsaea ensifolia,
L. odontolabia, Tectaria humbertiana, Tricho-
manes longilabiatum, and Xiphopteris sp. nov. 4.
Other species are restricted to narrow and frag-
mented biotopes in which it would be difficult or
impossible to delimit 800 m 2 plots that contain a
homogeneous vegetation. These include Blech-
num bakeri and B. ivohibense, which occur pref-
erentially at the bases of large rocks; Cyclosorus
interruptus, which grows in swampy areas; Ma-
crothelypteris torresiana and Odontosoria mel-
leri, which are found in open areas; and Elapho-
glossum petiolatum ssp. salicifolium, Lindsaea
odorata, Pityrogramma calomelanos, Sphaero-
stephanos arbuscula, and Sphenomeris chinensis,
which colonize rocky banks of rivers and streams.
In addition, three species and one variety (Lyco-
podiella caroliniana, Pityrogramma humbertii, P.
humbertii var. ?, and Schizaea pectinata) appear
to be restricted on Marojejy to xerophytic grass-
lands found above the upper limit of forests, be-
tween 2000 and 2132 m, a biotope that has a total
of only eight pteridophyte species and that was
not sampled quantitatively as part of the present
study. In all, only one species and two varieties
of pteridophytes known to occur rather frequently
in forest habitats on Marojejy were absent from
the study plots: Asplenium herpetopteris var. acu-
tipinnata, Athyrium scandicinum var. bipinnata,
and Nephrolepis tuberosa.

The shapes of the species-area curves (Figs. 2-
7A-D) indicate that the sample plots are both flo-
ristically homogeneous and representative of the
biotopes in which they were located. The curves
for the 18 plots are regular and give no indication
of intermediate plateaus, suggesting that the
boundaries of each plot fell entirely within a sin-
gle biotope. Each of the curves shows a clear as-
ymptote, except for PI 3 (Fig. 2-7D), which cov-
ered only 600 m 2 , and P8, P10, and R3 (Fig. 2-
7C), which had particularly rich pteridophyte flo-
ras, with about 60 species each. For pteridophytes,
the minimal plot size needed in the PN de Ma-
rojejy to establish a nearly complete sample of the
floristic composition thus appears to be between
600 and 1,000 m 2 , the precise size being a func-
tion of the level of species richness of the biotope
sampled. In practice it is often difficult to estab-
lish plots as large as 1,000 m 2 in which the veg-
etation is homogeneous, especially at higher alti-
tudes above 1500 m and on ridges. The average
value of 800 m 2 adopted for the present study thus

RAKOTONDRAINIBE: PTERIDOPHYTES

33

-a

T3

c

'C

-a

T3

c

(=

a

r2

oo oo

oo

oo

00

00

a. a.

oo

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c

00

h

o o

c

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c

c

03

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_o

03

03

XI

03

"oo ~

jO

00

00

-C

B3
g

X)

00

JU _cu

O

ju

E

g

u

=3 ^

ca

~

rS

03

03

T3

o -o

a>

~

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<L>

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u-

l

^

'e'e

00

'1

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7

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00

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c

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g g

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c x x x >-:

CL, PQ PL) W Dh

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PL)

v." ul ui ^ ^
CL, CL, Dh PJ PJ

Dh pq

X

PL)

pq

LD

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P 5 5 S 5

5j .3 5 s-

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a .g ^ su

R R v. 5?"

-q s 5

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."^ .*^ .*^ ."^ ."^ ."" -*^ 2 2 if.

k r r e 5 r '

5- S- S-

on =o t^
< < ^

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.-2

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s o o

R C R

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I s g

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2 ^
S 5.

2
g

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34

FIELDIANA: ZOOLOGY

u

U

PQ

C/3 W3

c/3 C/3

-^ -^

.* ^

c c

C C

03 03

03 03

X) X

X) X>

E g

E E

c 03

03 03

<U <U

<u <U

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w w

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5 ?

S 5 S

sill

a g *
s 3 C

las .

o, a, o, as

^ ^ c s

p* r* a n

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g.3

^ "a

j~ -a

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8 s %

c -a ^

b -a

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2 E

5

a ?u ^>

a a B

a a

2 5 S

a b o a

>s <n in in

%>

a a
a a
-a -a

co co

2 -a -a

J ^ (J u u

(NmONW00000\^fO\O
OOnOOOO- 'tNfNrO
--(SN(NN(SN(N(N(S

c-9

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E

appears to be an acceptable compromise (see also
Rakotondrainibe, 1989).

The stratified sampling method used here does
not make it possible to determine precise upper
and lower altitudinal limits for the floristic groups
of pteridophytes defined in the study. Continuous
sampling along an altitudinal transect would be
necessary, but this method was not compatible
with the logistical constraints of the fieldwork per-
formed or with the objectives of the multidisci-
plinary approach adopted, which were to evaluate
the overall biodiversity of a protected area in a
limited amount of time (45-60 days).

Vegetation Stages and Floristic Groups of
Pteridophytes on the Eastern Slope of the
Marojejy Massif

Humbert (1955) and Guillaumet et al. (1975)
recognized four principal vegetation types on the
Marojejy Massif: (1) moist evergreen forests of
low and middle altitude, from sea level to ca. 800
m; (2) montane forests, or moss forests with an
herbaceous lower layer (the latter term used by
Perrier de la Bathie, 1921), between ca. 800 and
1450 m; (3) dense sclerophyllous high-altitude
forests, or lichen forests, between about 1400-
1450 and 1850 m; and (4) montane thickets or
ericoid bushland, above about 1900 m. The phys-
iognomy and the flora of these four vegetation
types on Marojejy were described in detail by
Humbert (1955) and remain essentially unaltered
to this day, except in some areas around 400-550
m altitude, where the primary forest has given
way to a more open formation, often dominated
by a native bamboo species (Ochlandra capitata
E. G. Camus) or a Zingiberaceae (Aframomum an-
gustifolium K. Schum.).

The pattern of distribution of pteridophytes
generally coincides with that of the vegetation
types, although some exceptions can be noted at
both lower and upper ends of the altitudinal gra-
dient, where water or humidity can become a lim-
iting factor, as follows:

Between 400 and 850 m the physiognomy of
the vegetation changes progressively along the
eastern slope and shifts imperceptibly from low-
to middle-altitude forest. However, the pterido-
phyte groups A and B, whose limit occurs in this
same area, have very different floristic composi-
tions, as indicated by their respective positions on
the first two axes of the CA and their separation
at a high level in the HAC. Pteridophytes are par-

RAKOTONDRAINIBE: PTERIDOPHYTES

35

A Plots at low attitude (470-530 m)

-P1

-P2

P3

-P4

-R1

100 200 300 400 500 600 700 800
Area (m*)

B Plots at middle altitude (740-820 m)

P5

P6

P7

-R2

70

60

100 200 300 400 500 600 700 800
Area (m 2 )

C Plots at middle altitude (1290-1310 m)

P8

P10

-R3I

O

70

100 200300400500600 700 800
Area(m 2 )

D Plots at high altitude (1530-1920 m)

-P9

-P11

-P12

-P13

-P14

-P15

100 200 300 400 500 600 700 800
Area (m 2 )

Fig. 2-7. Accumulation curves of pteridophyte diversity in each plot within the PN de Marojejy. A, plots at low
altitude (470-530 m): B, plots at middle altitude (740-820 m); C, plots at middle altitude (1290-1310 m): D, plots
at high altitude (1530-1920 m).

ticularly sensitive to soil and atmospheric humid-
ity (Dzwonko & Kornas, 1994: Kornas. 1993;
Rakotondrainibe & Guillerm. 1990; van der
Werff. 1990; Young & Leon, 1991). At low alti-
tudes of 400-500 m. i.e.. below the zone in which
substantial condensation of fog takes place, the
presence of even a short dry season prevents the
most drought-intolerant species from growing,
such as Asplenium sandersonii, Blechnum atten-
uatum, Blotiella madagascariensis, Cochlidium
serrulatum, Ctenopteris devoluta, C. zenkeri, Cy-
athea bullata, and numerous species of Elapho-
glossum, Grammitis, Hymenophyllum, and Lind-

saea. These same taxa occur, however, toward
750-850 m. at least on the lower slopes and in
the valleys, where conditions favor their presence.
By contrast, at the other end of the altitudinal
spectrum, between 1500 and 2000 m. an abrupt
shift occurs at about 1800 m. from sclerophyllous
forest to montane thicket vegetation, whereas little
change occurs in the floristic composition of the
pteridophytes. The plots comprising group D are
separated into two subgroups, Dl and D2 (PI 2.
P13. P9 and PI 1. P14. P15, respectively) only
along axis IV of the CA. and even then at a rather
low level in the HAC. This relative homogeneity

36

FTELDIANA: ZOOLOGY

of the pteridophyte flora in such drastically dif-
ferent ecological conditions is surprising. The
species typical of sclerophyllous forests, in which
the canopy reaches 8-15 m in height, grow in a
shaded, continually moist forest environment,
whereas those in the montane thickets occur in a
low vegetation type that is subjected to marked
daily variations in temperature and insolation. The
latter species are, however, small in size, grow
among mosses that are saturated with water, and
form layers around branches and a dense carpet
on the ground. Thus, it is possible that water re-
serves in the substrate may allow species that are
sensitive to microclimatic variations to survive
and occur abundantly in the more buffered parts
of the sclerophyllous forest. Lack of data on the
amount of sunshine received on the summit of
Marojejy prevents the formulation of any formal
hypotheses. However, Guillaumet et al. (1975) in-
dicated that measurements made at 2050 m alti-
tude during the month of November (one of the
driest in the region) showed that the soil was per-
manently saturated with water at a depth of only
3 cm below the surface. If this observation ac-
curately reflects the situation on Marojejy
throughout the year, it would explain the relative
homogeneity of the pteridophyte groups above
1500 m altitude. Observations made in the equiv-
alent vegetation types in the Anjanaharibe-Sud
Massif, located at approximately the same latitude
but 35-40 km farther inland, support this idea.
There, contrary to what has been found on
Marojejy, the floristic composition of the pteri-
dophytes in montane thicket vegetation is statis-
tically different from that found in adjacent scler-
ophyllous forest (Rakotondrainibe & Raharimal-
ala, 1998). Observations in the field showed,
however, that during November 1996, the moss
layer in the montane thicket, which occurs on a
narrow ridge at 1950 m altitude, just below the
summit, dries out rapidly in the midday sunshine.
These observed differences confirm the idea that
pteridophytes are valuable indicators of microcli-
mate.

Species Richness

The biodiversity of three protected areas in
eastern Madagascar PN de Marojejy, PN
d'Andohahela, and RS d' Anjanaharibe-Sud) (Fig.
2-1) has now been studied using identical meth-
ods. Among these areas, Marojejy has by far the
richest pteridophyte flora, with 239 species, as

compared to 207 and 211 species, respectively,
for the other two areas (Rakotondrainibe, 1999;
Rakotondrainibe & Raharimalala, 1998), an ob-
servation that can be explained by climatic, bio-
logical, and geomorphological factors. The PN de
Marojejy is located only 45 km from the east
coast of Madagascar in a region with the highest
rainfall in the country (Donque, 1975). The east
slope of the massif is directly exposed to the trade
winds; annual rainfall at sea level in Antalaha av-
erages 2150 mm, and it is almost certainly higher
between 500 and 2132 m on Marojejy. Mean tem-
perature in Antalaha is 29.8C in the warmest
month and is 18.1C in the coolest month; thus it
has a warm, humid climate particularly favorable
for the growth of ferns. The Anjanaharibe-Sud
Massif has a similar climate, although it is most
likely somewhat less humid due to its more inland
location. The PN d'Andohahela, situated in ex-
treme southeastern Madagascar, just outside the
tropics, is in an area with a somewhat cooler and
drier climate (Donque, 1975; Rakotondrainibe,
1999). At both Marojejy and Andohahela, pri-
mary forest (including montane thickets) that has
been minimally affected by humans covers a wide
altitudinal belt, from 400 to 2000 m. By contrast,
at Anjanaharibe-Sud the lower limit of the native
forest does not extend below 800 m except for a
few patches near 600 m altitude. Also, PN de Ma-
rojejy and PN d'Andohahela are approximately
the same size (60,050 and 63,100 ha, respective-
ly), much larger than RS d' Anjanaharibe-Sud
(18,255 ha). Unlike the two other reserves, Ma-
rojejy is surrounded by several associated massifs,
and air flow in the numerous valleys and along
the slopes generates a multitude of microclimates
that can support greater levels of biodiversity.

In the three massifs studied to date, species
richness and species density peak in the mid-al-
titudinal level, between 750 and 1350 m, i.e., in
the zone where condensation from fog and oro-
graphic precipitation occur, where topography has
the most influence, and where the diversity of
habitats is thus greatest. This appears to be a
widespread phenomenon: in Panama, where the
highest peaks terminate at 3374 m, pteridophyte
richness is greatest between 500 and 1500 m (Lel-
linger, 1985); in South Africa the maximum rich-
ness occurs between 1000 and 1500 m on massifs
that reach to 3000 m, and in the high massifs of
East Africa pteridophyte richness is greatest be-
tween 1500 and 2000 m (Jacobsen & Jacobsen,
1989). On Mt. Kinabalu, in Borneo, which reach-

RAKOTONDRAINIBE: PTERIDOPHYTES

37

Fig. 2-8. Species richness of the 10 largest pterido-
phyte genera in the PN de Marojejy. RS d*Anjanaharibe-
Sud. and RNI d'Andohahela.

es to 4175 m altitude, maximum diversity was
observed at about 1500 m (Parris et al., 1992).

Figure 2-8 shows the number of taxa (species
and varieties) of the 10 most speciose pterido-
phyte genera in the Marojejy, Anjanaharibe-Sud,
and Andohahela reserves. The pteridophyte flora
of Marojejy is distinctive in being relatively poor
in members of the genus Asplenium (22 taxa vs.
26 and 30, respectively, for the other two areas)
and having more representatives of the genera
Grammitis (12 taxa vs. 9 and 7) and especially
Lindsaea (10 taxa vs. 4 and 2). The other genera
listed have approximately equivalent numbers of
taxa in the Marojejy and Anjanaharibe-Sud re-
serves, and in most cases there are fewer repre-
sentatives in PN d'Andohahela. It is not possible
at present to suggest a possible explanation for
these observed differences in floristic composi-
tion, but ecological and geographic factors surely
play a role. Mapping now under way to depict the
distributions of taxa with restricted ranges in
Madagascar and to assess their affinities with taxa
present in other floristic zones should, however,
make it possible to formulate hypotheses.

Floristic Distinctiveness

The fern flora of the Marojejy Massif has a
high level of endemism, but it also exhibits strong
floristic similarities with its closest neighbor, the
Anjanaharibe-Sud Massif. Table 2-8 shows the
coefficients of floristic similarity (Ps) (S0rensen,
1948) between the two areas, calculated on the

Table 2-8. Degree of similarity of the pteridophyte
flora of PN de Marojejy and RS d* Anjanaharibe-Sud by
altitudinal zone (Ps = S0rensen coefficient of similarity
calculated on the basis of the quantitative plot data).

Altitudinal zone (m)

Ps (%)"

740-2000

740-1000

1250-1350

1500-2000

74.2
66.0
59.1
61.3

* Ps = 200 X C/A+B. where A = species number in
the PN de Marojejy. B = species number in the RS
d' Anjanaharibe-Sud. and C = species number shared be-
tween the PN de Marojejy and RS d' Anjanaharibe-Sud.

basis of the quantitative plot data. Analysis of the
full set of plots located between 740 and 2000 m
(i.e., the range of altitudes at which primary forest
occurs at both sites) yields a very high coefficient
of similarity (Ps = 74.2%). Comparison of the
different altitudinal zones, however, shows the
strongest differences in the floristic composition
occurring in the middle altitude zone, between
1250 and 1350 m (Ps = 59.1%), which is char-
acterized by the presence of a closed, high tree
canopy. Homosporous pteridophytes, with their
very light diaspores that can travel over great dis-
tances, are the most likely pteridophyte taxa to
disperse between relatively isolated forest blocks
such as Marojejy and Anjanaharibe-Sud. How-
ever, the exchange of diaspores between the two
massifs may be impeded at middle altitudes,
where a closed forest canopy presents a barrier to
dispersal. This might explain in part the lower lev-
el of floristic similarity at these altitudes.

Figure 2-9 summarizes the chorology of the
239 species and varieties of pteridophytes ob-

6 7

6

B Marojejy (6)

H*T '{$k

^10

Q Marojejy + Anjanaharibe (7)
E3NE Madagascar (6)

1161

176

5 N Madagascar (10)

H Madagascar (76)

D Malagasy Region (18)

18

Wider distribution (116)

Fig. 2-9. Number of taxa (species and varieties) lo-
cally and regionally endemic among the 239 pterido-
phytes known from the eastern slopes of the Marojejy

Massif.

38

FIELDIANA: ZOOLOGY

served on the eastern slope of Marojejy. A total
of 6 taxa are endemic to the massif; 7 are known
only from Marojejy and Anjanaharibe-Sud; 6 are
restricted to the northeastern region of Madagas-
car, including these two reserves as well as the
Masoala Peninsula and the areas around Vohemar
and Mananara-Nord; and 10 taxa have distribu-
tions covering the northern part of Madagascar,
north of a line from Mananara-Nord to Antsohy.
The floristic distinctiveness of Marojejy in-
creases with altitude (Table 2-5). The number and
percentage of species endemic to Madagascar in-
crease regularly between 400 and 2000 m, and
four of the six species known only from the east-
ern slope of the massif are found in the highest
altitudinal zone. It is difficult to explain the ob-
served discrepancy between the strong floristic
similarity and the presence of several taxa endem-
ic to each of these two massifs, which are in close
geographical proximity and have comparable al-
titudinal ranges and geological features. It would
be interesting to observe the species diversity of
the corridor forests between the two reserves.

Conclusions

An assessment of the pattern of distribution of
pteridophyte taxa present on the eastern slope of
the Marojejy Massif was conducted using com-
plementary numerical methods. The results pro-
vide baseline data on the current species richness,
floristic composition, and endemism of the area.
Because the park is about to be opened to eco-
tourism, it is hoped that this study will be useful
to those responsible for the Marojejy/Anjanahar-
ibe-Sud Integrated Conservation and Develop-
ment Project in their effort to manage and con-
serve the biodiversity of these protected areas,
which are without doubt among the richest in
Madagascar.

Acknowledgments

I thank S. Goodman for the invitation to par-
ticipate in the fieldwork and to contribute a chap-
ter to this volume. I also thank B. Randriamam-
pionona for his technical assistance in the field,
S. Comtet for data entry, and N. d'Amico and A.
Pouron for collection curation. I am very grateful
to P. Lowry, G. Aymonin, and C. Sastre for fruit-

ful discussion. Special thanks go to P. Lowry for
translating the manuscript from the original
French version.

Literature Cited

Benzecri, J. P., and F. Benzecri. 1984. Pratique de
1' analyse des donnes. 1. Analyse des correspondances,
expose elementaire, 2 e ed. Bordas, Paris, 456 pp.

Bishop, L. E. 1978. Revision of the genus Cochlidium
(Grammitidaceae). American Fern Journal, 68(3): 76-
94.

Des Abbayes, H. 1961. Lichens recoltes a Madagascar
et a la Reunion (Mission H. Des Abbayes, 1956). I.
Introduction. II. Parmeliacees. Me'moires Institut
Scientifique de Madagascar, serie B, 10(2): 81-122.

Donque, G. 1975. Contribution geographique a l'etude
du climat de Madagascar. Nouvelle Imprimerie des
Arts Graphiques, Antananarivo, Madagascar, 477 pp.

Dzwonko Z., and J. Kornas. 1994. Patterns of species
richness and distribution of pteridophytes in Rwanda
(Central Africa): A numerical approach. Journal of
Biogeography, 21: 491-501.

Guillaumet, J.-L., J.-M. Betsch, C. Blanc, P. Morat,
and A. Peyrieras (sous la direction de R. Paulian).
1975. Etude des ecosystemes montagnards dans la re-
gion malgache. III. Le Marojezy. IV. L'ltremo et
l'lbity. Geomorphologie, climatologie, faune et More
(Campagne RCP 225, 1972-1973). Bulletin du Mu-
seum National d'Histoire Naturelle, serie 3, 309
(Ecologie generate 25): 29-67.

Holttum, R. E. 1974. Thelypteridaceae of Africa and
adjacent islands. Journal of South African Botany, 40:
(2) 123-128.

Humbert, H. 1955. Une merveille de la nature a Mad-
agascar: Premiere exploration du massif du Marojejy
et de ses satellites. Memoires Institut Scientifique de
Madagascar, serie B, 6: 1-210.

Jacobsen, W. B. G., and N. H. G. Jacobsen. 1989.
Comparison of the pteridophyte flora of southern and
eastern Africa, with special reference to high-altitude
species. Bulletin du Jardin Botanique National de Bel-
gique, 59: 261-317.

Koechlin, J., J.-L. Guillaumet, and P. Morat. 1974.
Flore et vegetation de Madagascar. J. Cramer Verlag,
Vaduz, 687 pp.

Kornas, J. 1993. The significance of historical factors
and ecological preference in the distribution of Afri-
can pteridophytes. Journal of Biogeography, 20: 281-
286.

Kramer, K. U. 1972. The Lindsaeoid ferns of the Old
World, IX. Africa and its islands. Bulletin du Jardin
Botanique National de Belgique, 42: 305-345.

Kramer, K. U., and P. S. Green. 1990. Pteridophytes
and gymnosperms. In Kubitzki K., ed., The Families
and Genera of Vascular Plants. Springer- Verlag, Ber-
lin, 404 pp.

Lebart, L., A. Morineau, and M. Piron. 1997. Statis-
tique Exploratoire Multidimentionnelle, 2 3 ed. Dunod,
Paris, 439 pp.

RAKOTONDRAINIBE: PTERIDOPHYTES

39

Lellinger, D. B. 1985. The distribution of Panama's
pteridophytes. Monographs in Systematic Botany.
Missouri Botanical Garden. St. Louis 10: 43-47.

Nicoll. M. E.. and O. Langrand. 1989. Madagascar:
Revue de la conservation et des aires protegees. World
Wide Fund for Nature. Gland, xvii + 374 pp.

Parris. B. S.. R. S. Beam an. and J. H. Beaman. 1992.
The plants of Mount Kinabalu 1 . Ferns and fern allies.
Royal Botanic Gardens. Kew. 165 pp.

Perrier de la Bathie, H. 1921. La vegetation malgache.
Annales de l'institut botanico-geologique colonial de
Marseille, ser. 3. 9: 1-268.

Pichi-Sermolli. R. E. G. 1977. Tentamen Pteridophy-
torum genera in taxonomicus ordinem redigendi.
Webbia. 31: 313-512.

Rakotondrainibe. F. 1989. Etude floristique. biologique.
phytogeographique et ecologique des fougeres et plan-
tes alliees de la foret d'Ambohitantely (foret tropicale
d'altitude). These de Doctorat. Universite de Lille.
304 pp.

. 1999. Pteridophytes of the eastern slope of the

Reserve Naturelle Integrate d*Andohahela. Madagas-
car: Distribution and floristic analysis, pp. 25-49. In
Goodman. S. M., ed., A floral and faunal inventory
of the Reserve Naturelle Integrale d'Andohahela.
Madagascar: With reference to elevational variation.
Fieldiana: Zoology, n.s.. 94: 1-297.

Rakotondrainibe. E. and J. L. Guillerm. 1990. Modele
de repartition horizontale des Pteridophytes le long
d'un gradient d'humidite lie a la topographie: le cas
d'une foret naturelle a Madagascar. Candollea. 45:
591-598.

Rakotondrainibe, E. and F Raharimalala. 1996. The
pteridophytes of the eastern slope of the Reserve Na-
turelle Integrale d'Andringitra. Madagascar, pp. 76-
82. In Goodman. S. M.. ed.. A floral and faunal in-
ventory of the eastern slopes of the Reserve Naturelle
Integrale d'Andringitra. Madagascar: With reference
to elevational variation. Fieldiana: Zoology, n.s.. 85:
1-319.

1998. The pteridophytes of the Reserve Spe-

With reference to elevational variation. Fieldiana: Zo-
ology, n.s.. 90: 1-246.

Rakotondrainibe, E. and N. Quansah. 1994. The di-
versity and originality of the pteridophytes of the for-
est of Manongarivo Special Reserve (north-west Mad-
agascar). Fern Gazette. 14(7): 259-268.

S afford. R. and W. Duckworth. Eds. 1990. A wildlife
survey of Marojejy Nature Reserve. Madagascar: Re-
port of the Cambridge Madagascar Rainforest Expe-
dition. 1988. Study Report no. 40. International Coun-
cil for Bird Preservation. Cambridge, 172 pp.

Schelpe. E. A. C. L. E. 1970. Pteridophyta. In Exell, A.
W, and E. Launert, eds.. Flora Zambesiaca. Crown
Agents, London, 254 pp.

Schelpe. E. A. C. L. E.. and M. A. Dintz. 1979. Pteri-
dophyta. In Mendes. E. J. ed.. Flora de Mozambique.
Junta de Investigacoes Cientiffcas do Ultramar. Centro
de Botanica, Lisboa. 257 pp.

Sorensen. T. 1948. A method of establishing groups of
equal amplitude in plant sociology based on similarity
of species content. Det Kongelige Danske Videnska-
bernes Selskab. Biologiske Skrifter. 5(4): 1-34.

Stefanovic. S.. E Rakotondrainibe. and E Badre.
1997. Selaginellacees. In Morat. P.. ed.. Flore de Mad-
agascar et des Comores. Paris. 68 pp.

Tardieu-Blot. M. L. 1951-1971. Les Pteridophytes. In
Humbert. H.. ed.. Flore de Madagascar et des Co-
mores. Museum National d'Histoire Naturelle Paris.

. 1955. Les Fougeres du massif du Marojejy et

Annexes, pp. 219-243. In Humbert. H. ed. Une merv-
eille de la nature a Madagascar: Premiere exploration
du massif du Marojejy et de ses satellites. Memoires
Institut Scientifique de Madagascar, serie B, 6: 1-210.
-. 1964. Les Pteridophytes. In A. Aubreville, ed..

ciale d*Anjanaharibe-Sud. Madagascar: Floristic anal-
ysis and altitudinal distribution, pp. 9-38. In Good-
man, S. M., ed.. A floral and faunal inventory of the
Reserve Speciale d'Anjanaharibe-Sud. Madagascar:

Flore du Gabon. Museum National d'Histoire Natu-
relle. Paris. 228 pp.

Tindale. M. D. 1965. A monography of the genus Las-
treopsis Ching. Contributions of the New South Wales
National Herbarium. 3(5): 249-339.

van der Werff. H. 1990. Ferns as indicators of vege-
tation types in the Galapagos archipelago. Mono-
graphs in Systematic Botanical Gardening, Missouri
Botanical Garden, St. Louis 32: 79-92.

Yolng. K. R.. and B. Leon. 1991. Diversity, ecology
and distribution of high-elevation pteridophytes within
Rio Abiseo National Park, north-central Peru. Fern
Gazette. 14(1): 26-39.

40

FIELDIANA: ZOOLOGY

Chapter 3

Structure and Floristic Composition of the Vegetation
of the Pare National de Marojejy, Madagascar

Nathalie Messmer, 1 Pierre Jules Rakotomalaza, 2 and Laurent Gautier 1

Abstract

A rapid biological inventory was conducted along an altitudinal transect in the Pare National
de Marojejy, Madagascar, between 4 October and 20 November 1996. Ten linear samples of
100 m were studied in five different elevational zones spanning 500 to 1950 m, and in a broad
assortment of forest types. Five hundred eighty-four species of vascular plants representing 73
families were censused in the linear samples. In addition, nine 0.1 ha temporary plots were
established in the lower four elevational zones. The data obtained provide quantitative and
qualitative information on the structure, biometrics, and floristics of the vegetation on the
Marojejy Massif.

Structural data revealed a progressive decrease in tree height and tree diameter as a function
of altitude. The density of trees >10 cm dbh was highest in mid-elevation plots. Basal area
was greatest at the lower altitudes. Dominant families changed gradually with altitude. Species
diversity was maximum at 1200 m. Although there was little similarity between samples, some
clusters were apparent and confirmed the trend observed in forest structure. The floristic and
structural differences between ridges and slopes at the same elevation were in general as im-
portant as the differences between one ridge sample and the slope sample at an altitude 400 m
higher.

Generally, the results of this study support Humbert's (1955, 1965) classification, but un-
derline the importance of topography in delimitation of forest types and the gradual transitions
from one type to another. Four hundred seventy-five fertile plants were collected, adding in-
formation to an already relatively well-known flora, based on the extensive collections of
Humbert (1955) and several other more recent collectors.

Resume

Une mission d'inventaire biologique rapide a ete menee aux mois d'octobre et novembre
1996 le long d'un gradient altitudinal dans le Pare National de Marojejy, Madagascar. Dix
releves lineaires de 100 m et neuf parcelles de 0.1 ha ont ete etudiees, fournissant ainsi des
donnees quantitatives et qualitatives, permettant 1'etude floristique, structurale et biometrique
de la vegetation du massif du Marojejy. Cinq cent quatre-vingts quatre especes de plantes
vasculaires reparties en 73 families ont ete recensees dans les dix releves lineaires.

Sur la base des donnees structurelles, on observe une diminution progressive de la hauteur
des arbres et de leur diametre en fonction de l'altitude. La densite des arbres >10 cm dbh
atteint sa valeur maximale dans les parcelles de moyenne altitude. L'aire basale est maximale

' Conservatoire et Jardin botaniques de la Ville de Geneve, Case Postale 60, CH-1292 Chambesy/Geneve, Swit-
zerland.

2 Missouri Botanical Garden, B.P. 3391, Antananarivo (101), Madagascar.

MESSMER ET AL.: VEGETATION 41

dans les parcelles de basse altitude. Les families dominantes changent progressivement avec
l'altitude. La diversite specifique atteint son maximum a 1200 m. Malgre une faible similarite
entre les releves, des regroupements sont possibles confirmant les tendances observees au ni-
veau de la structure. Les differences floristiques et structurales entre crete et versant a une
altitude donnee sont en general aussi importantes que celles observees entre un releve de crete
et le releve de pente 400 m plus haut.

De maniere generate, les resultats de cette etude confirment la classification de Humbert
(1955, 1965), mais soulignent l'importance de la topographie dans la delimitation des types de
vegetation, ainsi que la nature graduelle du passage d'un type de vegetation a un autre. Quatre
cent soixante quinze echantillons de plantes fertiles ont ete recoltes, s'ajoutant a ceux de Hum-
bert (1955), ainsi qu'a ceux d'autres recolteurs, completant ainsi petit a petit la connaissance
de la biodiversite floristique du Pare National de Marojejy.

Introduction

The remarkable flora of the Marojejy Massif
was first encountered by Henri Humbert during
his exploration of the northern mountains of Mad-
agascar, which also included the other mountains
ringing the Andapa basin and the Tsaratanana
Massif (Koechlin et al., 1974). Humbert visited
Marojejy on several occasions and considered it
one of the most beautiful and impressive massifs
of Madagascar; in a 1955 monograph he de-
scribed it as "Une merveille de la nature a Mad-
agascar" [A wonder of nature in Madagascar].
Humbert collected almost 2,500 specimens in the
Marojejy area (ca. 2,100 from the actual reserve),
which contributed significantly to establishing the
floristic diversity of this mountain. However, his
account was published as a series of notes and in
no way is a comprehensive floristic treatment of
the massif. Humbert (1955) also proposed a clas-
sification of the Marojejy forests, but this was
based on general floristic observations and per-
sonal knowledge rather than on quantitative sur-
veys.

Subsequently numerous other botanists made
important collections on Marojejy, and an im-
pressive list of plants and herbarium specimens
will soon be available (Miller et al., in prep.).
However, prior to our work, no quantitative or
structural sampling of the vegetation had been
done on the massif. The biological inventory of
the Pare National (PN) de Marojejy, conducted in
1996, provided an opportunity to fill this gap.

Humid forests are usually divided into altitu-
dinally segregated vegetational belts based on flo-
ristic components and estimated quantitative char-
acteristics (e.g., diameter at breast height [dbh],
height). Our specific objectives were (1) to pro-
vide a quantitative floristic account of the various

study sites, (2) to compare the floristic composi-
tion of different habitats and habitats at different
elevations on the massif, and (3) to document
vegetational changes along an elevation gradient.

Methods

Vegetation surveys in the tropics have long suf-
fered from the fact that the classic phytosociolog-
ical approaches developed for use in temperate
regions are not applicable because of the high spe-
cies diversity and complexity of tropical ecosys-
tems. A number of techniques have been devised
to overcome this obstacle almost as many tech-
niques as there are groups of researchers working
in this field. Whereas the high degree of floristic
and structural variation between different types of
tropical forests may warrant the use of different
approaches, the lack of a common survey tech-
nique hampers comparisons among studies.

For tropical forests, 1 ha plots have been wide-
ly used, but they have a number of disadvantages:
generally they take into account only individuals
with dbh >10 cm, which leaves out herbs, shrubs,
lianas, and most regenerating (i.e., young) tree
species. They also do not provide precise results
on vegetation structure. Establishing the plot itself
is a time-consuming operation and generally dam-
ages part of the understory. In short, for several
reasons the use of these plots in rapid inventory
assessments is not ideal.

The methods used in rapid biodiversity assess-
ment surveys should be simple but sufficiently
comprehensive to allow characterization of criti-
cal aspects of the local flora and comparisons with
other studies. To address these requirements, Gau-
tier et al. (1994) have proposed a method that

42

FIELDIANA: ZOOLOGY

combines sampling vegetation above 100 points
regularly spaced along a line and sampling in
small temporary plots. The linear sample provides
an estimation of cover of the main species, which
is a direct indication of their importance in the
community. For tree species, the linear sample
can be compared with the Importance Value Index
of classic 1 ha plots. In addition, the linear sample
gives detailed results on vegetation structure (ver-
tical and horizontal distribution of vegetation).
The small temporary plots are located along the
line of the linear sample. They allow rapid mea-
surement of main biometric parameters such as
number of trees and basal areas. The combined
technique of linear sampling and small plot sam-
pling is relatively easily implemented in the field,
even in difficult environments with a thick under-
story, such as in secondary re growth. It is also
suitable for steep and rapidly changing reliefs,
where large surfaces for 1 ha plots often are not
available. The time needed to establish and survey
one linear sample and the associated temporary
plot, including the collection of voucher speci-
mens, is ca. 2.5-3 days in a dense forest with a
relatively unknown flora.

Study Sites and Sampling Procedure

Fieldwork was conducted at five sites along an
elevational gradient in the PN de Marojejy rang-
ing from near 500 m to 1950 m. The study zones
were centered on the five camps (at 450, 775,
1225, 1625, and 1875 m) and sampling was con-
ducted within an elevational swath of 75 m. The
precise localities of each camp and their geore-
ferenced coordinates are given in Chapter 1 (pp.
5-6). Two habitats were studied in each of these
sites, one on a slope (on a gradient intermediate
between ridgetop and valley bottom) and one on
a ridge (along the crest of a ridge). These two
kinds of sites were chosen to better represent po-
tential microhabitat variation within each eleva-
tional zone. The only exception was at 500 m,
where, because of topographical features, flat pla-
teaus rather than distinct ridges were found. In the
highest elevational zone, only linear sampling was
done.

An effort was made to choose floristically ho-
mogeneous sampling sites. The forest along the
elevational gradient, however, had been partially
disturbed by human activities. Indeed, in the first
study zone, at 450 m, there were areas of dis-
turbed forest, and selective logging was not rare.

In this elevational zone we chose sites for linear
samples and plots in the least disturbed areas. In
the second elevational zone, at 775 m, some evi-
dence of selective cutting was also present. At
higher altitudes, selective extraction of some tree
species was noted, but there was no extensive per-
turbation.

Linear Samples

The linear sampling technique involved record-
ing contacts of vegetation with a vertical line ex-
tending from 100 points set along a baseline at
ground level. It is generally recommended that the
length of the baseline be around 10 times the
height of the canopy, but in our case, topograph-
ical features forced us to work along a 100 m line.
Measurements were made at 100 points, 1 m
apart. The height of contact and species encoun-
tered above each of the 100 points were recorded.
To do so, we erected a vertical 8 m pole con-
structed out of 5 segments each 1.6 m long, the
entirety demarcated in 20 cm increments. The
heights at which all plants trees, shrubs, lianas,
herbaceous plants, and seedlings came in con-
tact with this pole were recorded for each of the
sampled points. Heights of contacts above 8 m
were estimated. When the vertical line passed
through dense foliage, the lower and upper points
of contact of this foliage mass were recorded.
When plants could be unambiguously identified
in the field, their scientific names were recorded
on the data sheet. If the identity of a plant was in
doubt, a voucher specimen was collected and the
species was referred to as a "morphospecies" on
the data sheet. These morphospecies were sequen-
tially numbered or named independently for each
altitudinal zone. If a voucher specimen could be
unambiguously identified as identical to another
specimen collected at the same site, it received
the same binomial or morphospecies number and
then the other specimen was discarded. Fertile in-
dividuals were also systematically collected.

The sterile material collected in the plots and
along the linear samples was identified in the field
at least to family level. After the material was
dried, global morphospecies determinations were
made in the herbarium of the Pare Botanique et
Zoologique de Tsimbazaza (TAN), Antananarivo,
and the herbarium of the Direction des Ressources
Forestieres et Piscicoles (= FO.FI.FA) (TEF), An-
tananarivo. In some cases binomial names could
be assigned to these sterile specimens on the basis

MESSMER ET AL.: VEGETATION

43

of a fertile specimen represented in the general
collections.

Plots

Temporary plots of 0.1 ha (100 X 10 m or 50
X 20 m) were marked, with the base of the linear
sample used as the median line of each plot. Trees
rooted within the plots and having a dbh >10 cm
were recorded and their dbh was measured. Plots
were not established in the highest elevational
zone (1800-1950 m), which encompassed the
ecotone between the upper lichen forest and high-
elevation thicket, because in the lower portion of
this zone the dwarf trees had less than the re-
quired dbh (>10 cm).

General Collecting

Within each elevational zone, fertile specimens
were collected from time to time outside the linear
samples and plots but within the elevational swath
of the site for additional floristic information. A
maximum of seven specimens of each fertile in-
dividual were collected. The first specimen of
each plant was deposited in the herbarium of TAN
or TEF; the other specimens were sent to the her-
baria of the Missouri Botanical Garden, St. Louis
(MO), the Conservatoire et Jardin botaniques de
la Ville de Geneve (G), and the Museum National
d'Histoire Naturelle de Paris (P). For difficult
groups, a specimen was also sent to a specialist
for identification. Remaining duplicate specimens
were designated to be sent to the Royal Botanic
Gardens, Kew (K), and to the Agricultural Uni-
versity of Wageningen (WAG).

Data Analysis

Linear Samples

Vegetation Profile The data collected in the
linear sample give a comprehensive view of a
cross-section of the forest along a vertical plane
above the baseline. Plotting all contacts on a
graph with the length of the baseline as the jc-axis
and the height of the contacts as the y-axis gives
an informative visual representation of the distri-
bution of the vegetation. For a correct represen-
tation, the scales of both axes should be identical.
In this representation, it is also possible to assign

specific symbols for the main species to give an
idea of their distribution in the profile.

Vertical Distribution of Vegetation The
contacts with vegetation can be separated verti-
cally into height intervals. For each interval, the
occurrence of vegetation above each of the 100
points of the baseline is summed to give a percent
value of cover. The height of the interval can be
even, or it can follow any other rule. We chose a
first interval of 0-2 m for herbaceous and under-
growth vegetation, followed by a geometric two-
fold progression: 0-2 m, 2-4 m, 4-8 m, 8-16 m,
16-32 m, >32 m. This height-array is often se-
lected for tropical forests and has proved to be
very informative. The data are then plotted on a
vertical histogram where the length of each bar
represents the cover of each interval. An interest-
ing analysis of the shape of such histograms is
presented in Chatelain (1996).

Species Cover Species cover is a direct in-
dication of species' importance in the community.
If the number of points is 100, as in our case,
species cover can be directly inferred from the
number of points above which the species is re-
corded, and expressed as a percentage. For tree
species, a decreasing cover value generally re-
flects the ranking that can be obtained through
more complicated procedures, such as calculating
the Importance Value Index, at least for the prev-
alent species (Gautier et al., 1994). The number
of individuals involved in these contacts is also
derived from the data sheet and indicated beside
the percentage cover value. Especially in our
study plots, where topographical features limited
the length of the baseline to 100 m, big tree spe-
cies can exhibit percentage cover values that
might be heavily dependent on this limitation and
reach a nonrepresentative value, too low or too
high. The latter artifact becomes obvious when a
high percentage cover is associated with a small
number of individuals, such as one or two.

Species Cover per Height Interval The
same calculation can also be made after the in-
formation is first separated into height intervals as
defined above. The percentage cover of the spe-
cies in each interval is directly derived from the
sum of the occurrences of the species above the
100 points of the baseline. It should be noted that
for any species, the sum of the percentage cover
in all intervals is generally higher than the global
percentage cover because a species is often re-
corded in several intervals above a single point.

Species-Number of Points Accumulation
Curve The cumulative number of species en-

44

FIELDIANA: ZOOLOGY

countered along the sample can be plotted with
respect to the number of points. The accumulation
curve is similar to the classic species-area accu-
mulation curves of surface plots and follows the
same rules. Its shape and values are an indication
of the homogeneity and the floristic richness of
the sample. If the sample is homogeneous, a reg-
ular decrease in slope toward a horizontal asymp-
tote is expected.

The Shannon-Weaver Diversity Index, H'
Species diversity can be quantified by means of a
diversity index. For this survey we used the Shan-
non-Weaver diversity index, H' (Shannon &
Weaver, 1949). This frequently used index takes
into account not only the presence or absence of
a species, but also its numerical importance. In
this case the quantitative parameter used to cal-
culate the index was the number of individuals
censused along the lines of the linear samples.
This diversity index increases with diversity. For
two samples with an identical number of species,
the index is lower if a few species are dominant
than if the majority of species have a similar den-
sity.

The Horn Similarity Index, Ro The Horn
similarity index, derived by comparing the Shan-
non-Weaver diversity indices of a number of
plots, gives a useful measure of affinities in com-
munity composition (Brower et al., 1990). Its val-
ue ranges from (no similarity) to 100%.

additional information on a community's struc-
ture. Diameter classes are defined in 10 cm incre-
ments. The data are plotted as a histogram of the
number of individuals according to increasing di-
ameters. An inverted-J-shaped histogram is ex-
pected for mature forests.

Results

Description of the Samples

The results of the sampling are presented in de-
tail in Appendix 3-1, along with a profile high-
lighting the species with a cover >10% in any
height interval, the vertical distribution of the
vegetation (% cover), the points-species accu-
mulation curve, the number of stems 5:10 cm dbh,
basal area, the number of species, and the Shan-
non diversity index, as well as a list of the species
recorded, with cover values separated into height
intervals. The samples themselves are briefly de-
scribed below.

500 m, Slope

Appendix 3- la

Temporary Plots

For the estimation of biometric parameters such
as density and basal area of the community, it has
been shown that when these values are plotted
against an increasing sample area, the values at
first fluctuate widely, and an accurate value can
be reached only with plots of at least 0.16 ha
(Gautier et al., 1994). Because of time constraints,
it was possible to sample only 0. 1 ha in each site,
and this limitation should be remembered when
one interprets the values obtained in this study.

Density The density defined here is the num-
ber of trees with a dbh >10 cm.

Basal Area Basal area is the sum of the area
of the cross-section of the trunk at 1.3 m of each
tree with a dbh >10 cm. It is directly calculated
from the diameter measurements of the trees
(Centre Technique Forestier Tropical [CTFT],
1989).

Distribution of dbh The distribution of num-
ber of trees according to diameter classes gives

The first sample was chosen in a rather dense
lowland rain forest. All height intervals had a cov-
er value >60%, with the exception of the >32 m
stratum, which included three emergent species
(Albizia sp.l;* Brochoneura acuminata (Lam.)
Warb.;t cf. Deinbollia sp.l). The maximum cover
value (93%) occurred at the 16-32 m interval,
with an important contribution from Chrysophyl-
lum boivinianum (Pierre) Baehni, which also ac-
counted for half of the cover of the 8-16 m in-
terval. The 4-8 m interval was dominated by On-
costemum sp.l, Diospyros sp.2, and the epiphytic
fern Asplenium nidus L. Oncostemum sp.l was
also important in the 2-4 m and 0-2 m intervals,
together with two species of Rubiaceae and sap-
lings of numerous species.

* Attribution of genera to families is given in Appen-
dix 3-2.

t Authorities for species are given at the first mention
of the species in the text.

MESSMER ET AL.: VEGETATION

45

490 m, Plateau

Appendix 3- lb

At the same altitude, this second sample also
was a typical lowland rain forest. The emergent
stratum was denser and comprised six species, in-
cluding Brochoneura acuminata and Canarium
boivinii Engl. The 8-16 m and 16-32 m intervals
were dominated by Chrysophyllum boivinianum,
along with Lauraceae sp.l, Mallotus sp.l. and
Macaranga sp.3, the two latter species also dom-
inating the 4-8 m interval. Tabernaemontana sp.l
and the tree fern Cyathea decrescens Mett. shared
an important cover in the 4-8 m and 2-4 m in-
tervals. The lower interval was broadly dominated
by Poaceae sp. 1 . The profile gives the impression
that the vegetation was rather loose, which was
confirmed by the relatively low cover of the 16-
32 m interval. Although no stumps were visible
in the surroundings, this zone may have been se-
lectively logged in the past.

800 m, Slope

Appendix 3-lc

At this altitude the forest was slightly lower
than in the 500 m zone, and there were no emer-
gent species above 32 m. The profile shows a
rather discontinuous canopy around 25-30 m that
is not reflected in the high cover value of the 16-
32 m interval (74%) because of the contribution
of some medium-sized trees below. However,
there is a clear gap in the right-hand side of the
profile, and the presence of secondary species
(e.g., Aframomum angustifolium (Sonn.) K.
Schum., Ravenala madagascariensis Sonn.) in the
undergrowth indicated a past disturbance. In the
other parts of the profile, important species in-
cluded Albizia sp.l, Cryptocarya sp.3, Chryso-
phyllum boivinianum, and Sloanea rhodantha
(Baker) Capuron in the 16-32 m interval, and
Deuteromallotus sp.l and Symphonia sp.4 in the
4-8 m and 8-16 m intervals.

800 m. Ridge

Appendix 3- Id

As in the preceding sample, there were no
emergent species. The profile shows a very dense
vegetation with a closed canopy dominated by
Uapaca sp.2, Canarium boivinii, and two species
of Lauraceae in the 16-32 m interval. In the 8-

16 m interval, Uapaca sp.2 was also important,
together with Mammea sp.2 and Eugenia sp.l.
Two species of Pandanus, one Dypsis sp., and one
Cyathea sp. reached important cover values on the
forest floor.

1200 m, Slope

Appendix 3-le

The 1200 m slope sample had a continuous
canopy that was included in the 16-32 m interval,
except for a portion of the right-hand side of the
profile, which was more open. Main species in-
cluded Ephippiandra sp.2. an unidentified Rubi-
aceae. and Brachylaena sp.l. In the 8-16 m in-
terval the vegetation was also relatively dense,
with two species of Mammea and Eugenia sp.8.
The 4-8 m interval included Dypsis sp.4 and the
tree fern Cyathea sp.3. The 2-4 m interval has a
relatively poor cover value; vegetation included
Tabernaemontana sp.l. which was also important
in the 500 m plateau sample. In turn, the lower
interval reaches a high cover value with an un-
identified pteridophyte species that accounted for
half of the cover, and an unidentified Acantha-
ceae. With 103 species this sample exhibits the
highest diversity value of the survey, with a still
increasing species accumulation curve.

1200 m, Ridge

Appendix 3- If

The ridge sample at 1200 m showed a much
smaller forest: no contacts were recorded above
16 m. The forest was very dense below 10 m,
with some emergents. including Cryptocarya sp.6.
another unidentified Lauraceae. and Eugenia sp.4.
The 4-8 m interval included the same species,
together with Canthium sp.2 and Elaeodendron
sp.l. In the 2-4 m interval there were numerous
species with little species cover, except for Dypsis
sp.4, which is worth mentioning here. The lower
interval consisted mainly of lianescent bamboos,
with two Nastus species accounting for more than
half of the cover. The species accumulation curve
displays a strong increase at the middle of the
baseline corresponding to a slight change in slope
along the ridge. The number of species encoun-
tered (97) was also very high.

46

FIELDIANA: ZOOLOGY

Table 3-1. Distribution of diameters, density, and basal area for nine 0.1 ha plots in four different elevational
zones in the PN de Marojejy.

Total no.

Altitude

Habitat

Number of trees per dbh i

'in cm) classes

of trees
with dbh
i >10 cm

Basal
area

(m)

10-20

20-30

30-40

40-50

50-60

60-70

70-80

80-90 90-10C

(m 2 )

500

Slope

30

6

6

5

3

5

1

3

59

6.6

490

Plateau

34

12

6

2

1

1

2

1

59

3.8

800

Slope

46

19

4

3

1

1

2

76

4.4

800

Plateau

45

17

8

6

3

3

1

1

1

85

6.5

800

Ridge

90

33

11

4

1

3

1

143

6.7

1200

Slope

61

14

6

1

1

1

2

86

4.0

1200

Ridge

170

24

7

4

205

4.9

1600

Slope

116

40

6

1

163

4.6

1600

Ridge

7

2

1

10

0.3

1600 m. Slope

Appendix 3-lg

Forest structure here was rather similar to the
preceding sample, with almost no contacts above
16 m, a few emergents above 10 m (Eugenia sp.8
and Eugenia sp.4), and no clear stratification be-
low. The 4-8 m interval was widely dominated
by Dypsis sp.4 and Polyscias sp.5. The two lower
intervals share a relatively low cover value
(<70%), with numerous species.

1950 m, Ridge

Appendix 3-lj

The vegetation at the highest altitude sampled
was a dense ericoid thicket, with most contacts
occurring below 2 m. The species with the highest
cover value were Nastus sp.4, another unidentified
Poaceae, and a species of Weinmannia. Among
the shrubs species were also several Ericaceae
(Erica spp.; Agauria salicifolia (Comra. ex Lam.)
Hook. f. ex Oliv.) and Myrsinaceae. The herba-
ceous species included numerous Asteraceae.

1600 m, Ridge

Appendix 3-lh

The ridge sample at this altitude showed a very
dense dwarf forest structure with no stratification.
Most of the contacts were below 8 m. Main spe-
cies included Calophyllum sp.6, Nastus sp.2, and
Dypsis sp.6. Asteraceae and Poaceae were rich in
species and accounted for an important cover val-
ue. The number of species reached 98.

1850 m, Slope

Appendix 3-li

The structure of this sample was very similar
to the preceding one, with a slightly denser 4-8
m interval that was dominated by Calophyllum
sp.6 and Cassinopsis sp.4. The lower interval also
included Dichaetanthera sp.8, an unidentified
fern, and some lianescent bamboos (Nastus spp.).

Forest Structure

Table 3-1 summarizes the biometrical data ex-
tracted from the 0.1 ha plots nested in the linear
surveys (number of trees per diameter classes,
density, basal area). An additional plot was sam-
pled at 800 m elevation on a plateau. No plots
were established in the linear surveys at 1 850 and
1950 m because of the lack of trees >10 cm dbh.

The density of trees >:10 cm dbh ranged from
100 to 2,050 individuals per hectare. The lowest
value was found on the ridge at 1600 m. The
highest values occurred on the ridges at 800 and
1200 m and on the slope at 1600 m. They were
mainly due to the contributions of the low-diam-
eter vegetation classes (<30 cm dbh).

Figure 3-1 graphically expresses the distribu-
tion of dbh in 10-cm-increment classes for each
plot. The dwarf structure of the 1600 m ridge sites
is obvious, and for a clear interpretation of lig-
neous vegetation structure the smaller diameters
should also have been sampled. All histograms fit

MESSMER ET AL.: VEGETATION

47

500 m
slope

1600m
ridge

10-20 (cm) ]

20-30 (cm)

30-40 (cm)

40-50 (cm) I

50-60 (cm)
:H60-70(cm)

70-80 (cm)
S80-90 (cm) !

90-100 (cm)

i J

Fig. 3-1. Distribution of the individuals in 10-cm-increment dbh classes for nine 0.1 ha plots in four elevation
zones in the PN de Marojejy. Within each plot, the order of the bars corresponds to the key read from top to
bottom.

a classic inverted J curve, but some differences
are noteworthy: the two 500 m plots share a rel-
atively low number of individuals with dbh <20
cm. For >30 cm dbh. the plateau plot has low
figures, which are reflected in the basal area value.
The slope plot in turn has remarkably steady val-
ues in the >20 cm dbh classes: compared to the
other histograms, the 20-30 cm class is very low
and the >50 cm classes are high. The 1200 m
ridge and 1600 m slope plots display a compres-
sion of the histogram toward the low values of
diameter. The 1 200 m ridge plot has two big trees
in the 80-90 cm class, which is quite rare at this
altitude. In the 800 m slope plot, there is a clear
gap in the 50-60 cm class.

Basal area ranges from 3 to 67 mTha. The low-
est value again occurred on the ridge at 1600 m.
The highest values do not match the high values
for density except for the 800 m ridge plot. The
high basal area values of the 500 m slope and the
800 m plateau plots are mainly due to a relatively
high number of trees with a dbh >30 cm.

The structural parameters extracted from the
ten linear surveys are shown in Figures 3-2 (pro-
files) and 3-3 (cover per height interval histo-
grams).

A few general observations on forest structure
from the linear survey data were made in the brief
descriptions of the samples, but combining all the
data in a single figure adds new comparative in-
formation. Both samples at 500 m are quite sim-
ilar, with the exception of the 16-32 m height in-

terval, which has more cover on the slope site.
This finding is related to the high value of the
>50 cm dbh classes and the high basal area men-
tioned earlier. At 800 m the slope site has lower
cover values than the ridge site (especially below
16 m): this finding is again linked to the basal
area and distribution of dbh classes.

From 800 m upward a gradual decrease in veg-
etation height is apparent. Also, within each alti-
tude zone, vegetation on the slope site was sys-
tematically taller than vegetation on the ridge site.
Furthermore, both the profiles and the cover per
height interval histograms show that the forest
structure of the ridge site is very similar to that
of the slope site at the next higher altitude inter-
val.

Forest Composition

Along the elevational transect. 3,011 angio-
sperm individuals representing 584 morphospe-
cies in 73 families were censused (Appendices 3-
2 and 3-3). In addition. 475 specimens of fertile
plants were collected during the general collec-
tions, representing 64 families of angiosperms
(Appendix 3-4). The list of the species present in
each linear survey, along with their cover values,
is given in Appendix 3-1 and was briefly com-
mented on in the description of the samples.

The cover values of plant families along the ten
linear samples is given in Appendix 3-3. For the

48

FIELDIANA: ZOOLOGY

b) Plateau at 500 m

10 20 30 40 50 60 70 80 90 100
Length (m)

10 20 30 40 50 60 70
Length (m)

35

30

_25

20

lb 15
J 10

1 5

c) Slope at 800 m

r.

b

%cS^^w^4a

35

30

? 25

r2o

15
I 10

d) Ridge at 800 m

10 20 30 40 50 60
Length (m)

70 80 90 100

30 40 50 60
Length (m)

70 80 90 100

35

30

_25

E.20

I, 15

S 10
X
5

e) Slope at 1200 m

f) Ridge at 1200 m

70 80 90 100

35
30

? 25

r2o

X 10
5

g) Slope at 1600 m

^S^S^tera

35

30

25

^20

5 15

X 10

5

h) Ridge at 1600 m

20 30 40 50 60
Length (m)

70 80 90 100

35

30

-25

20

1 5

i) Slope at 1850 m

10 20 30 40 50 60 70 80 90 100

35

30

-25

r 2 o

5 IS

3= 10
5

j) Ridge at 1950 m

30 40 50 60
Length (m)

10 20 30 40 50 60
Length (m)

70 80 90 100

Fig. 3-2. Vegetation profile of the ten 100 m linear samples conducted in five elevation zones in the PN de
Marojejy. Data represent height of vegetation contacts measured or projected above the 100 points of the baseline
linear sample.

main families the results are also expressed graph-
ically in Figure 3-4. Some plant families were
clearly restricted to low altitudes (Myristicaceae,
Fabaceae, Annonaceae, Moraceae, Burseraceae,
Apocynaceae) or, if represented at all altitudes,
showed a clear preference for low elevation
(Ebenaceae, Sapotaceae, Euphorbiaceae, Rubi-
aceae). Lauraceae were already dominant in the
500 m plots, but their cover values continued to
increase up to 1200 m. Clusiaceae were respon-
sible for an important percentage of cover at all
altitudes. Poaceae were present in all samples and

reached important values in the highest plots as
well as in the 500 m plateau plot. Myrsinaceae
and Elaeocarpaceae also occurred at all altitudes,
but with rather low cover values. Pandanaceae,
Monimiaceae, Acanthaceae, Myrtaceae, Areca-
ceae, Melastomataceae, and Flacourtiaceae were
likewise recorded at all altitudes, but their highest
cover value occurred in the 1200-1600 m forests.
Finally, Araliaceae, Asteraceae, Cunoniaceae,
Balsaminaceae, and Ericaceae were seen mainly
above 1200 m, and these families displayed an
increasing affinity for high altitude.

MESSMER ET AL.: VEGETATION

49

c) Slope

It Mill ill

32

1

_ 16-32

] -i

5 8-16

!

M

5 *

2-4

H so

0-2

10 20 30 40 50 60 70 80 90 100
Cover percentage

e) Slope a

1200 m

>32

16-32

j

8-16

PJgO

3 4-8

X

2-4

M&' 4

SB 4 *

0-2

Hit

10 20 30 40 50 60 70 80 90 100
Cover percentage

10 20 30 40 50 60 70 80 90 100
Cover percentage

i) Slope

.It

18511 i

>32

16-32

8-16

1

4-8

I'M

2-4

71

10 50 60 70 80 90 100
r percentage

10 20 30 40 50 60 70 80 90 100
Cover percentage

10 20 30 40 50 60 70 80 90 100
Cover percentage

h) Ridge at 1600 m

>32

1

16-32

8-16

i

4-8

2-4

H74

0-2

flHxi

10 20 30 40 50 60 70 SO 90 100
Cover percentage

j) Ridge at 1950 m

>32

16-32

8-16

4-8

2-4

I 8

0-2

10 20 30 40 50 60 70 80 90 100
Cover percentage

Fig. 3-3. Vertical distribution of vegetation (cover percentage in six height classes) for the ten 100 m linear
samples surveyed in five elevational zones in the PN de Marojejy.

The number of species recorded in each linear
survey is shown in Table 3-2, together with the
Shannon-Weaver diversity index and the species/
individual ratio. The highest numbers of species
recorded occurred in the 1200 m samples. It
should be noted that the species accumulation
curves (Appendix 3-1) do not approach a satura-
tion value at the end of 100 m of the baseline in
all samples, with (perhaps) the exception of the
two samples at highest elevation.

The values of the Shannon- Weaver diversity in-
dex (H') for species vary between 1.56 and 1.80
(for families, between 1.15 and 1.37) in the sam-

pled sites (Table 3-2). The degree of diversity
quantified by this index is rather constant along
the altitudinal transect, but the plots at low alti-
tude and the 1950 m ridge have lower values.
Taking into account what we said earlier regard-
ing the species accumulation curves, this fact
could be attributed to insufficient sampling at the
two 500 m samples, but the 1950 m forest could
in fact be less diverse.

The Horn similarity index values for species
and families are presented in Table 3-3. The value
calculated at the species or morphospecies level
between ridge and slope sites of each elevational

50

FIELDIANA: ZOOLOGY

C. O 03 i
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S O. O CQ

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pil

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Cu 00 J u

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MESSMER ET AL.: VEGETATION

51

Table 3-2. Values of Shannon-Weaver diversity in-
dex for each linear transect in the PN de Marojejy.

Species/

Altitude
(m)

Habitat

No.
of
Spe-
cies

Shannon-Weaver
diversity index

Species Families

No.
of indi-
viduals

ratio

500
490

Slope
Plateau

66
66

1.56
1.60

1.29
1.25

0.29
0.30

800
800

Slope
Ridge

70
78

1.67
1.73

1.37
1.35

0.36
0.27

1200
1200

Slope
Ridge

103
97

1.74
1.76

1.33
1.26

0.33
0.27

1600
1600

Slope
Ridge

82
98

1.67
1.80

1.28
1.29

0.26

0.24

1850
1950

Slope
Ridge

80

73

1.72
1.64

1.31
1.15

0.24
0.31

zone shows that important differences exist be-
tween the microhabitats within each elevational
zone as well as between the altitudinal belts. None
of these values shows more than 0.28 similarity.
The index values at the species level would prob-
ably have been higher if the number of samples
within each elevational zone had been increased.
However, time was always a limiting factor in sur-

veying the numerous requisite samples. Even if
the values of specific similarity are very low, the
differences between these values still reveal sam-
ple clusters, which will be discussed later.

If the values for the 500 m samples, for which
no ridge sample was available, are placed aside,
the mean value of the similarity index for slope
and ridge samples at the same altitude is 0.182.
The mean value of the similarity index for a ridge
sample and the slope sample at the next higher
altitude is almost the same (0.187), whereas the
mean value for a slope sample and the next higher
slope sample is much lower (0.044). These figures
further support the relation between the forests on
ridges and slope plots at the next higher altitude
and the intricateness of the forest types, as was
noted earlier in the discussion of forest structure.

When comparisons are made at the family level
the values of the Horn index are much higher, rang-
ing from 0.35 to 0.88. The distribution of these
values supports the trends observed in Figure 3-4.

Discussion

In his classic account of the vegetation types of
Madagascar, Humbert (1965) described the east-

Table 3-3. Values of the Horn similarity index at the morphospecies level (above the diagonal in italics) and at
the family level (below the diagonal) between the linear transects in the PN de Marojejy.

Slope Plateau Slope Ridge Slope Ridge Slope
500 m 490 m 800 m 800 m 1200 m 1200 m 1600 m

Ridge Slope Ridge
1600 m 1850 m 1950 m

Slope
500 m
Plateau
490 m
Slope
800 m
Ridge
800 m
Slope
1200 m
Ridge
1200 m
Slope
1600 m
Ridge
1600 m
Slope
1850 m
Ridge
1950 m

1.000

0.277

0.185

0.052

0.040

0.019

0.000

0.000

0.000

0.000

0.673

1.000

0.274

0.091

0.035

0.020

0.004

0.000

0.000

0.012

0.721

0.795

1.000

0.121

0.040

0.015

0.008

0.018

0.004

0.000

0.614

0.603

0.664

1.000

0.090

0.085

0.109

0.086

0.024

0.051

0.523

0.648

0.687

0.682

1.000

0.188

0.177

0.080

0.055

0.102

0.662

0.589

0.617

0.816

0.757

1.000

0.242

0.095

0.032

0.006

0.45 1

0.530

0.505

0.748

0.775

0.774

1.000

0.193

0.091

0.037

0.503

0.461

0.434

0.681

0.778

0.608

0.752

1.000

0.133

0.062

0.480

0.429

0.445

0.636

0.748

0.666

0.743

0.877

1.000

0.254

0.456

0.354

0.364

0.502

0.703

0.560

0.639

0.769

0.837

1.000

Note: Bold entries represent medium index values (0.7-0.8 for families and 0.1-0.2 for species); bold underscored
entries represent high index values (>0.8 for families and ^0.2 for species).

52

FIELDIANA: ZOOLOGY

ern Malagasy humid forest and cited certain gen-
era (or families) as abundant. In our study, Chry-
sophyllum, Symphonia, and the Lauraceae were
indeed abundant, but Dilobeia, also cited as char-
acteristic by Koechlin et al. (1974), was never re-
corded in this study. However, Humbert (1955)
recorded Dilobeia thouarsii Roem. and Schult.,
but on Mt. de Beondroka, not on the main massif
of Marojejy, and Miller (pers. coramun.) noted
that this species is common at some sites on Ma-
rojejy and Beondroka. To our knowledge, this
species has never been recorded on the Tsaratan-
ana or Manongarivo Massifs. It should also be
mentioned that the genus Podocarpus was not
found during our survey of the PN de Marojejy.
Humbert (1955) did not observe this genus on the
main slopes of Marojejy, although he found it in
the foothills of the massif, for example on Mont
d'Ambatosoratra, on Mont de Beondroka, and in
the Ambatoharanana Valley (Laubenfels, 1972).
At other surveyed mountain sites, such as the
eastern slopes of the PN d'Andringitra, Podocar-
pus was one of the dominant taxa at 1600 m
(Lewis et al., 1996). It is also present in the Re-
serve Speciale de Manongarivo (pers. observ.)
and on the Tsaratanana Massif, but absent in the
PN d'Andohahela (Rakotomalaza & Messmer,
1999) and RS dTvohibe (pers. observ.). Given the
uneven distribution of Podocarpus, it should no
longer be considered one of the characteristic gen-
era of eastern Malagasy forest from the littoral to
high-altitude forests, as stated by Koechlin et al.
(1974).

The eastern Malagasy humid forest contains a
number of distinct vegetational communities that
vary as a function of altitude. On the basis of the
classification presented by Humbert (1955), which
he later generalized to the whole eastern slope
(1965), the eastern rain forest on the Marojejy
Massif runs from the lower limit of the reserve to
about 800 m. (The various classifications of these
vegetation formations used by different research-
ers are given in Table 3-4.) Humbert's mid-ele-
vation rain forest falls between 800 m and 1450
m; his lichen forest generally falls between 1450
m and 1800 m; and finally high-elevation thicket
and grassland form a mosaic above 1800 m up to
the summit at 2137 m. The results of our survey
will now be compared with this classification.

In the lower portion of the lowland humid for-
est, the presence of Mauloutchia humblotii (H.
Perrier) Capuron, Chrysophyllum boivinianum,
Shane a rhodantha, and Canarium sp. (not all
these species were recorded in the samples, but

they were found in the surrounding forest) follows
closely the characteristic floristic features of this
formation as proposed by Humbert (1965). Mau-
loutchia humblotii is restricted to the lowest stud-
ied level, 500 m, and was not found at the 800 m
site; the upper limit of this species is probably
about 500-600 m. The upper limit of Chryso-
phyllum boivinianum stands at 1000 m.

The four samples around 500 m and 800 m al-
titude share a common structure, especially when
the distribution of dbh is considered (Fig. 3-1).
The profiles (Fig. 3-2) and vertical distribution of
vegetation (Fig. 3-3) nevertheless show that the
forests at 500 m are of a slightly taller stature.
However, these forest structures fit well with
Humbert's (1955) description of the eastern rain
forest, with a canopy between 25 and 30 m high.
The floristic composition of the slope sample at
800 m is an extension of that in the lowland hu-
mid forest as recorded at about 500 m (similarity
index > 0.15; Table 3-3). The species composi-
tion of these three samples shows very little sim-
ilarity (<0.05) with the samples at or above 1200
m. The ridge plot at 800 m has relatively low
affinities with all other plots and may represent a
transition in floristic composition although still re-
lated to the lowland rain forest in terms of struc-
ture.

The slope sample at 1200 m, with its 18-25 m
canopy and a less important medium stratum, cor-
responds typically to the mid-elevation forests of
Humbert. The presence of Ephippiandra and
Tambourissa (Monimiaceae), two altitudinal in-
dicator genera, supports this conclusion (Hum-
bert, 1965; White, 1986). Lauraceae is also one
of the dominant families, especially above 800 m,
and this observation holds for most of the Mala-
gasy humid forest (Lewis et al., 1996; Rabevoh-
itra et al., 1996; Lowry et al., 1997; Rakotomalaza
& Messmer, 1999).

In turn, the 1200 m ridge sample structure has
more affinities with the description of the lichen
forest (6-12 m canopy, intermingled with the
shrub stratum), like the 1600 m slope sample. The
change in forest composition between 1 200 m and
1600 m is exemplified by the disappearance of
Canarium sp. and Sloanea rhodantha, both spe-
cies characteristic of lowland rain forest and mid-
altitude rain forest up to 1200 m that were ob-
served only in the slope habitats and not on the
ridges at this altitude. Moreover, the appearance
of bamboos and an increase in cover values for
Clusiaceae such as Symphonia spp. support the
observation. Indeed, bamboos and Symphonia are

MESSMER ET AL.: VEGETATION

53

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54

FIELDIANA: ZOOLOGY

indicators of high-altitude and exposed formations
such as ridges (Koechlin et al., 1974). The tran-
sition between mid-elevation rain forest and li-
chen forest would then occur between 1200 m and
1600 m (Table 3-3). The distinctly exposed posi-
tion of the ridge sample at 1 200 m, which shares
0.24 similarity with the 1600 m slope sample,
would explain why this site can be categorized
with the lichen forest formation.

The height of vegetation in the 1600 m ridge
sample is still lower, like the 1850 slope sample.
Both show a unique, dense stratum of vegetation
with a maximum height of about 6-7 m. Structur-
ally, these formations could be considered as be-
longing to the small-sized type of lichen forest. The
small trees with reduced leaves and small-diameter,
twisting trunks are also diagnostic of this formation
(Koechlin et al., 1974). The floristic composition
is also characteristic, as indicated by the presence
of Alberta sp., Symphonia sp., Weinmannia spp.,
several species of Asteraceae, and a wide assort-
ment of Myrtaceae, Lauraceae, and Araliaceae
(Schefflera spp. and Polyscias spp.).

Above 1900 m the summital areas are subjected
to high variations in humidity and temperature
(Koechlin et al., 1974), conferring on the vege-
tation its particular aspect. The forest is reduced
to high-elevation thicket, in which the ridge sam-
ple at 1950 m was established. It consists of a
very low and dense thicket with Ericaceae woody
plant genera Erica and Agauria, as well as Cryp-
tocarya (Lauraceae). It corresponds to the ericoid
thicket of Humbert (1965). Between the patches
of this ericoid scrub, herbaceous high-altitude
grassland rich in many Asteraceae, such as Heli-
chrysum, Vernonia, and Senecio, dominates the
landscape. The mosaic of these types of vegeta-
tion formations in the highest elevational zone es-
sentially follows variations in topography, with
high-elevation thicket on ridges and in valley bot-
toms and grassland on slopes.

A progressive change in vegetation was ob-
served along the elevational transect in the PN de
Marojejy. The family cover values in the samples
(Fig. 3-4) show a clear shift in composition with
altitude. However, as indicated by the Horn sim-
ilarity index (Table 3-3), this change is gradual,
and there is considerable overlap of families along
the elevational transect. The high values for the
Horn similarity index attest that the family taxo-
nomic level is a too large an entity with which to
characterize the different elevational vegetation
zones. Forest classification should be based not on
families but on species or morphospecies. Nev-

ertheless, some families are largely characteristic
of a given altitude, especially when their impor-
tance value is taken into account.

The shifts from one type of vegetation forma-
tion to another were gradual but readily apparent.
Furthermore, above 800 m floristic and structural
similarities were noted between a ridge at a given
elevation and the slope of the next highest zone
(e.g., between the ridge at 1200 m and the slope
sample at 1600 m). Hence, the distribution of veg-
etation types does not correlate with altitude
alone. Moreover, as already noted by Humbert
(1955), it is unlikely that the divisions will occur
at the same elevation everywhere on the Marojejy
Massif or along the same latitudinal gradient in
Madagascar. Topography plays an important role
in the scheme of demarcations between each al-
titudinal vegetation formation.

On a larger scale, a progressive downward
shifting of elevational zonation toward the south
of Madagascar may exist. We have already noted
the downward shift of the botanical classification
limits along the altitudinal gradient in the PN
d'Andohahela (Rakotomalaza & Messmer, 1999).
The results of the three 1.0 ha permanent plots
established in the PN de Ranomafana (Lowry et
al., 1997) also suggest that the altitudinal bound-
ary between lowland rain forest and mid-altitude
rain forest stands at 600 m instead of the classic
800 m. The plot at 1200 m, moreover, is different
from the ones at 600 and 950 m, which are sim-
ilar. The boundary, then, between mid-altitude
rain forest and lichen forest should fall some-
where between 950 and 1200 m. These observa-
tions support a compression of the altitudinal veg-
etation zones along a latitudinal gradient from
north to south.

To compare the biometric parameters deter-
mined during our study of the PN de Marojejy
with other sites on Madagascar or elsewhere in
the tropical world, we multiplied the 0. 1 ha plots
by 10 to bring them to the standard area of 1.0
ha. A comparison with other eastern humid forest
sites in Madagascar shows overall similarities be-
tween these forests (Table 3-5). However, com-
parisons between Madagascar and tropical forests
elsewhere in the world show that the number of
trees with stems >10 cm dbh and basal area val-
ues are much higher in the PN de Marojejy than
at sites in South America, Africa, and Asia (see
Rakotomalaza & Messmer, 1999, for a review).

Regarding diversity, the fact that the number of
species was highest at 1200 m could also be inter-
preted as an artifact of the length of the baseline

MESSMER ET AL.: VEGETATION

55

Table 3-5. Comparative plot data from other parts of Madagascar.

Number

Shannon-

Shannon-

of trees

Weaver

Weaver

with dbh

Basal

diversity

diversity

Plot size

Altitude

>10 cm

area (m 2 )

index

index

Sites

(ha)

(m)

per ha

per ha

families

species

Miaranony. Ranomafana (Lowry et al..

1

600

769

49.9

1997)

Vatoharanana. Ranomafana (Lowry et al..

1

950

660

35.0

1997)

Vohiparara. Ranomafana (Lowrv et al..

1

1200

1.092

25.6

1997)

Manombo (Rabevohitra et al., 1996)

1

80

787

Analalava-Marovony (Dumetz. 1993)

1

50

1.200

4.90

1

50

840

4.60

1

50

940

4.70

1

50

1.280

5.00

Manatantely (Dumetz. 1993)

1

250

900

3.90

Andohahela ( Rakotomalaza & Messmer,

1

440

739

34.1

1.32

1.76

1999)

1

840

880

43.2

1.34

1.86

1

1140

1.216

43.8

1.29

1.77

1

1550

675

63.8

1.18

1.49

1

1875

1.365

65.9

1.21

1.52

Marojejy. biological inventory of 1996

0.1

490

590

38.0

1.25

1.60

0.1

500

590

66.0

1.29

1.56

0.1

800

860

65.0

0.1

800

760

44.0

1.37

1.67

0.1

800

1.430

67.0

1.35

1.73

0.1

1200

880

40.0

1.33

1.74

0.1

1200

2.050

49.0

1.26

1.76

0.1

1600

1.640

46.0

1.28

1.67

0.1

1600

100

3.0

1.29

1.80

1850

1.31

1.72

1950

1.15

1.64

Note: The Shannon-Weaver diversity index values were calculated from the linear transect in the PN de Marojejy.

of the surveys (100 m), which is probably too short
to allow adequate representation of the species, es-
pecially for the bigger trees in the lower-altitude
samples. However, in 1 ha plots in the PN
d' Andohahela, the same observation has been
made for angiosperms (Rakotomalaza & Messmer.
1999) as well as for pteridophytes in the RS
d'Anjanaharibe-Sud (Rakotondrainibe & Rahari-
malala. 1998). and thus the trend could be real. The
species Shannon- Weaver diversity index value we
measured in the PN de Marojejy is much lower
than the ones calculated for other overseas tropical
regions (reviewed by Rakotomalaza & Messmer.
1999). These values are nevertheless comparable
to the ones calculated from the results obtained
along the elevational transect in the PN
d' Andohahela (Table 3-5). but the ratios of the
number of species to the number of individuals are
much higher in the PN de Marojejy. However, the

meaning of these low values is not yet elucidated,
and the sample size might be implicated.

Because two different methods were used in the
PN de Marojejy and the PN d' Andohahela, and
owing to the insufficient amount of quantitative
data along the latitudinal gradient in the eastern
forests of the island, it would be premature to
draw meaningful conclusions about clinal varia-
tion in species richness associated with latitudinal
variation. This possibility, however, should be
considered when subsequent data become avail-
able from the rain forests of Madagascar.

Conclusions

The field methods used during the 1996 inven-
tory of the PN de Marojejy provided good quan-

56

FTELDIANA: ZOOLOGY

titative measures and graphic representations of
the structure, fioristic composition, and variability
of the vegetation along the elevational gradient.
Regarding structure, the linear survey gives valu-
able information on forest structure even for
dwarf forests, where trees >10 cm dbh are rare
(1600 m ridge, 1800 m slope, and 1950 m ridge
samples).

The linear sampling method has been used only
in a few tropical forests (but see Chatelain, 1996),
and comparisons with data obtained by other
means from other sites are limited. More use of
this method should be encouraged, because of its
efficiency during rapid inventories under difficult
conditions and the useful results it provides.

To allow a comparison of biometric parameters
such as basal area, density, and the like, tradition-
al plots are added to the linear sampling method.
However, these plot data would be of greater val-
ue if their area is increased to 0.2 ha, as recom-
mended by Gautier et al. (1994), and if the dbh
lower limit is reduced to 2.5 cm instead of 10 cm
in half of the plot.

The sampling method we used does not give a
full account of the diversity of the richest sites,
and problems in application of the diversity index
still have to be solved. However, the compromise
between the time devoted to each sample (ca. 2.5
days) and the data gathered renders the method
very valuable for rapid assessment surveys.

Comparisons would have been easier if we
would have applied the 0.1 ha transects (ten con-
tinuous transects, each 50 m X 2 m), as proposed
by Gentry (1982, 1988). Indeed, these 0.1 ha
"Gentry" transects have been applied in the trop-
ics at more sites than any other rapid assessment
fioristic evaluation technique. We did not, how-
ever, use this inventory method because it is often
described more as a complement to 1.0 ha plots
than as a survey method by itself (Phillips & Ra-
ven, 1996), aimed as it is toward capturing max-
imal diversity rather than accurate structural and
fioristic measurements. Moreover, a 0.1 ha "Gen-
try" transect probably traverses an area of ap-
proximately 2 or 3 ha (Phillips & Raven, 1996).
To locate a homogeneous parcel of forest meeting
this area requirement on the higher slopes of a
mountain such as Marojejy would not be simple.
Our aim was to describe two different habitats,
slopes and ridges, in a restricted altitudinal inter-
val. Thus, even if the 0.1 ha transect method is
regarded as one of the favorite methods for fio-
ristic inventories, it was not adequate for our sur-
vey objectives.

The quantitative survey we made in the PN de
Marojejy adds new data to that already recorded
by the numerous collectors on the Marojejy Mas-
sif area during earlier botanical expeditions. The
structural data add information about the eleva-
tional gradient in the northeastern forest of Mad-
agascar. We have referred to Humbert's classifi-
cation system in discussing our results obtained
in the PN de Marojejy because it has a more pre-
cise basis than many other earlier schemes and
because more recent classification systems are
largely based on that of Humbert.

Generally, the structure and fioristic composi-
tion of the forests along the altitudinal gradient
agrees with Humbert's (1955, 1965) visual de-
scriptions. The vegetation types he noted were
readily encountered in our samples, but some
shifts in altitude between slope and ridge vege-
tation types emphasized the influence of topog-
raphy. Furthermore, the changes in vegetation
types were gradual and the altitudinal gradient
surveyed in this study encompassed an ecocline
rather than clear-cut vegetation communities.
Similar surveys in the Malagasy rain forests
would help elucidate the relationship between al-
titudinal botanical zonation and latitudinal varia-
tion.

Acknowledgments

We thank Steve Goodman for inviting us to
take part in the biological inventory of the PN de
Marojejy. Many thanks to Desire Ravelonarivo,
who assisted us with much enthusiasm. The late
Jeanine Raharilala helped extensively with plant
identifications, and we are grateful to the staff of
the Pare Botanique et Zoologique de Tsimbazaza
for allowing access to the collection. We are
grateful to the Missouri Botanical Garden and the
Conservatoire et Jardin botaniques de la Ville de
Geneve for agreeing to have us participate to this
inventory, and we thank ANGAP and DEF for
arranging permits for our fieldwork. We are also
grateful to Steve Goodman for the detailed com-
ments and advice he gave. We thank James Miller
for the access to a draft of his work in preparation.
Finally, we thank an anonymous reviewer for use-
ful comments on the manuscript.

MESSMER ET AL.: VEGETATION

57

Literature Cited

Brower, J. E., J. H. Zar, and C. N. Von Ende. 1990.
Field and Laboratory Methods for General Ecology.
3rd ed. Brown Publishers, Dubuque, Iowa, 237 pp.

Centre Technique Forestier Tropical (CTFT). 1989.
Memento du forestier: Techniques rurales en Afrique.
Ministere de la Cooperation et du Developpement
(eds), Paris, 1266 pp.

Chatelain, C. 1996. Possibility's duplication de
l'imagerie satellitaire a haute resolution pour 1' etude
des transformations de la vegetation en Cote dTvoire
forestiere. These de doctorat, Universite de Geneve,
Departement de botanique et de biologie vegetale, Ge-
neve.

Cornet, A., and J.-L. Guillaumet. 1976. Divisions flor-
istiques et etages de vegetation a Madagascar. Cahier
de l'ORSTOM (serie Biologie), 11: 35-42.

Dumetz, N. 1993. Mise en evidence de l'heterogeneite
floristique et structurale de la foret orientale de basse
altitude a Madagascar. These de doctorat, Universite
de Paris VI, Paris.

Du Puy, D. J., and J. Moat. 1996. A refined classifi-
cation of the primary vegetation of Madagascar based
on the underlying geology: Using GIS to map its dis-
tribution and to assess its conservation status, pp.
205-318. In Lourenco, W. R., ed. Biogeographie de
Madagascar. Editions de TORSTOM, Paris.

Gautier, L., C. Chatelain, and R. Spichiger. 1994.
Presentation of a releve method for vegetation studies
based on fine-scale satellite imagery, pp. 1339-1350.
In Seyani, J. H., and A. C. Chikuni, eds. Proceedings
of the XIII Plenary Meeting of the AETFAT, Malawi,
vol. 2.

Gentry, A. H. 1982. Patterns of neotropical plant spe-
cies diversity, pp. 1-84. In Hecht, M. K., B. Wallace,
and G. T Prance, eds. Evolutionary Biology, vol. 15.
Plenum Press, New York.

. 1988. Changes in plant community diversity

and floristic composition on environmental and geo-
graphical gradients. Annals of the Missouri Botanical
Garden, 75: 1-34.

Humbert, H. 1955. Une merveille de la nature a Mad-
agascar: Premiere exploration botanique du massif du
Marojejy et de ses satellites. Memoires de lTnstitut
Scientifique de Madagascar, serie B, 6: 1-271.

. 1965. Description des types de vegetation, pp.

46-78. In Humbert, H., and G. Cours Darne, eds. No-
tice de la carte Madagascar. Extrait des travaux de la
section scientifique et technique de lTnstitut Francais
de Pondichery, hors serie no. 6.

Koechlin, J., J.-L. Guillaumet, and P. Morat. 1974.
Flore et Vegetation de Madagascar. Cramer, Vaduz.
Liechtenstein, 687 pp.

Laubenfels (de), D. J. 1972. Podocarpacees. Flore de
Madagascar et des Comores, 18: 1-22.

Lewis, B. A., P. B. Phillipson, M. Andrianarisata, G.

Rahajasoa, P. J. Rakotomalaza, M. Randriambolo-
lona, and J. F McDonagh. 1996. A study of the bo-
tanical structure, composition, and diversity of the east-
ern slopes of the Reserve Naturelle Inte'grale
d'Andringitra, Madagascar, pp. 24-72. In Goodman, S.
M., ed. A floral and faunal inventory of the eastern
slopes of the Reserve Naturelle Integrate d'Andringitra,
Madagascar: With reference to elevational variation.
Fieldiana: Zoology, n.s., no. 85.

Lowry, P. P., G. E. Schatz, and P. B. Phillipson. 1997.
The classification of natural and anthropogenic vege-
tation in Madagascar, pp. 93-123. In Goodman, S. M.,
and B. D. Patterson, eds. Natural Change and Human
Impact in Madagascar. Smithsonian Institution Press,
Washington, D.C.

Miller, J. S., et al. In prep. A checklist of the vascular
plants of Marojejy.

Perrier de la Bathie, H. 1921. La vegetation malgache.
Annales de lTnstitut Botanico-geologique Colonial de
Marseille (serie 3), 9: 1-226.

Phillips, O. L., and P. H. Raven. 1996. A strategy for
sampling neotropical forests, pp. 141-165. In Gibson,
A. C., ed. Neotropical Biodiversity and Conservation.
University of California Press, Berkeley and Los An-
geles.

Rabevohitra. R.. P. P. Lowry II, G. E. Schatz, H. Ran-
drianjafy, and N. Razafindrianilana. 1996. Assess-
ment of plant diversity and conservation importance
of east coast low elevation Malagasy rain forests: Rap-
port sur le projet. Centre National de la Recherche
Appliquee au Developpement Rural, Madagascar. De-
partement de Recherches Forestieres et Piscicoles,
Madagascar, and Biodiversity Support Program, Mis-
souri Botanical Garden, St. Louis.

Rakotomalaza, P. J., and N. Messmer. 1999. Structure
and floristic composition of the vegetation of the Re-
serve Naturelle Integrale d'Andohahela. Madagascar,
pp. 51-96. In Goodman, S. M., ed. A floral and faunal
inventory of the Reserve Naturelle Integrale
d'Andohahela, Madagascar: with reference to eleva-
tional variation. Fieldiana: Zoology, n.s., no. 94.

Rakotondrainibe, F, and E Raharimalala. 1998. The
pteridophytes of the Reserve Speciale d'Anjanaharibe-
Sud, Madagascar: Floristic analysis and altitudinal dis-
tribution, pp. 17-38. In Goodman, S. M., ed. A floral
and faunal inventory of the Reserve Speciale
d' Anjanaharibe-Sud, Madagascar: with reference to ele-
vational variation. Fieldiana: Zoology, n.s., 90.

Shannon, C. E., and W Weaver. 1949. The Mathe-
matical Theory of Communication. University of Il-
linois Press, Urbana, 117 pp.

White, F 1986. La vegetation de l'Afrique: Memoire
accompagnant la carte de vegetation de 1' Afrique
UNESCO/AETFAT/UNSO. Recherches sur les res-
sources naturelles XX. ORSTOM-UNESCO. 384 pp.

58

FIELDIANA: ZOOLOGY

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69

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FIELDIANA: ZOOLOGY

3*
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MESSMER ET AL.: VEGETATION

73

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m =

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FIELDIANA: ZOOLOGY

Appendix 3-3. Plant Families Censused in the Ten Linear Samples in Five Elevational
Zones in PN de Marojejy, with Their Cover Value (%)

490 m

500 m

Pla-

800 m

800 m

1200 m

1200 m

1600 m

1600 m

1850 m

1950 m

Family

Slope

teau

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Ridge

Slope

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Ridge Slope

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Acanthaceae

3

5

31

7

2

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2

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1

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5

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10

13

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5

39

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1

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3

2

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4

7

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19

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9

36

11

24

6

Arecaceae

14

3

1

24

50

40

79

17

14

4

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5

11

22

17

34

23

26

Balsaminaceae

7

7

14

16

Begoniaceae

3

2

Bignoniaceae

9

2

Burseraceae

9

10

3

22

6

Canellaceae

2

Celastraceae

2

1

3

Chloranthaceae

1

Clusiaceae

20

13

29

39

31

28

31

63

44

13

Convolvulaceae

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4

1

2

2

9

15

16

Cyatheaceae

29

1

Cyperaceae

3

2

6

10

Dilleniaceae

8

Ebenaceae

23

11

14

4

7

5

7

4

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8

19

1

2

9

12

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5

18

28

Erythroxylaceae

14

4

4

5

8

1

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20

62

57

58

9

30

3

14

4

4

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16

1

26

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14

4

19

28

25

9

20

11

Icacinaceae

2

1

4

8

24

4

Lauraceae

22

32

40

49

15

47

11

3

7

Liliaceae

1

1

Linaceae

4

Loganiaceae

4

3

6

Loranthaceae

5

1

3

Maranthaceae

1

Melanophyllaceae

1

4

Melastomataceae

3

1

3

5

19

21

7

18

31

13

Meliaceae

4

4

1

Menispermaceae

4

Monimiaceae

9

3

4

29

8

3

1

2

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3

14

19

5

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10

1

Myristicaceae

13

15

Myrsinaceae

47

1

8

14

14

11

9

7

4

13

Myrtaceae

12

13

5

28

12

42

37

26

13

9

Oleaceae

1

4

7

1

2

Orchidaceae

1

1

3

1

1

3

Pandanaceae

2

56

2

14

13

1

3

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1

6

4

2

Pittosporaceae

3

2

10

Poaceae

1

42

2

23

9

47

13

36

36

66

Pteridophyta excl. Cyatheaceae

38

38

44

5

88

20

50

4

17

31

MESSMER ET AL.: VEGETATION

95

Appendix 3-3. Continued

490 m
500 m Pla- 800 m 800 m 1200 m 1200 m 1600 m 1600 m 1850 m 1950 m
Family Slope teau Slope Ridge Slope Ridge Slope Ridge Slope Ridge

Rhizophoraceae

Rubiaceae 66 26 27 20 68 38 40 34 21

Rutaceae 4 112 1 1

Sapindaceae

Sapotaceae

Smilacaceae

Sterculiaceae

Strelitziaceae

Thymeleaceae

Tiliaceae

Urticaceae

Verbenaceae 1 113

Viscaceae 3 1

Xyridaceae 1

Zingiberaceae 7

4

66

26

27

20

68

38

4

1

1

2

1

18

3

5

4

6

65

49

27

2

4

9

2

3

4

8

1

3

5
3

2
12

1

1
6

3

2

96 FIELDIANA: ZOOLOGY

Appendix 3-4. Fertile Plant Specimens Collected During the General Collections
Along the Altitudinal Gradient in PN de Marojejy

Family

Genus and species

Collector*
and number

Altitude (m)t

Acanthaceae

Acanthaceae

Acanthaceae

Acanthaceae

Acanthaceae

Acanthaceae

Acanthaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Annonaceae

Apiaceae

Apocynaceae

Apocynaceae

Apocynaceae

Aquifoliaceae

Aquifoliaceae

Aquifoliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Araliaceae

Arecaceae

Asclepiadaceae

Asclepiadaceae?

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

cf. Rue I Ha sp.

Campnosperma sp.

Campnosperma sp.

Campnosperma sp.

Protorhus sp.

Protorhus sp.

Protorhus sp.

Protorhus sp.

Protorhus sp.

Sorindeia madagascariensis DC.

cf. Isolona sp.

Isolona sp.

Isolona sp.

Isolona sp.

Monanthotaxis sp.

Polyalthia emarginata Diels

Polyalthia emarginata Diels

Polyalthia emarginata Diels

Polyalthia sp.

Xylopia sp.

Enteromorpha sp.

Landolphia sp.

Tabernaemontana eusepala Aug. DC.

Tabernaemontana sp.

Ilex mitis (L.) Radlk.

Ilex mitis (L.) Radlk.

Ilex mitis (L.) Radlk.

Polyscias sp.

Polyscias sp.

Polyscias sp.

Polyscias sp.

Polyscias sp.

Polyscias sp.

Schefflera sp.

Schefflera sp.

Schefflera sp.

Dypsis sp.

Apodocephala sp.

Brachylaena merana (Baker) Humbert

Helichrysum adhaerens (DC.) R. Vig. & Humbert

Helichrysum adhaerens (DC.) R. Vig. & Humbert

Helichrysum sp.

Helichrysum sp.

Helichrysum sp

Helichrysum sp

Helichrysum sp

Helichrysum sp

Helichrysum sp

Helichrysum sp

Helichrysum sp

PJ 746

450 75

NM 257

775 75

NM 334

1625 75

NM 413

1875 75

NM 414

1875 75

PJ 776

775 75

PJ 895

1625 75

NM 314

1225 75

PJ 800

1225 75

PJ 871

1625 75

NM 235

450 75

NM 305

1225 75

PJ728

450 75

PJ752

450 75

PJ 806

1225 75

NM 238

450 75

NM 263

775 75

NM 215

450 75

NM 227

450 75

PJ 761

775 75

PJ 827

1225 75

NM 281

1225 75

PJ 836

1225 75

PJ 859

1625 75

PJ 725

450 75

NM 312

1225 75

PJ 902

1875 75

NM 223

450 75

PJ 748

450 75

PJ 820

1225 75

NM 306

1225 75

NM 350

1625 75

PJ 849

1625 75

NM 385

1875 75

NM 408

1875 75

PJ 726

450 75

PJ 757

775 75

PJ 787

1225 75

PJ944

1875 75

PJ798

1225 75

PJ 874

1625 75

PJ938

1875 75

NM 344

1625 75

NM 211

450 75

PJ919

1875 75

PJ 914

1875 75

PJ 768

775 75

NM 310

1225 75

NM 384

1875 75

PJ 925

1875 75

NM 337

1625 75

NM 343

1625 75

NM 375

1875 75

NM 382

1875 75

NM 383

1875 75

NM 422

1875 75

NM429

1875 75

PJ 855

1625 75

PJ924

1875 75

MESSMER ET AL.: VEGETATION

97

Appendix 3-4. Continued

Family

Genus and species

Collector*
and number

Altitude (m)t

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Asteraceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Balsaminaceae

Begoniaceae

Bignoniaceae

Bignoniaceae

Bignoniaceae

Bignoniaceae

Burseraceae

Burseraceae

Burseraceae

Burseraceae

Canellaceae

Celastraceae

Celastraceae

Chloranthaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Helichrysum sp.
Helichrysum sp.
Rochonia sp.
Senecio sp.
Senecio sp.
Senecio sp.
Senecio sp.
Senecio sp.
Senecio sp.
Senecio sp.
Senecio sp.
Vernonia sp.
Vernonia sp.
Vernonia sp.
Vernonia sp.
Vernonia sp.

Impatiens sp.
Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Impatiens sp.

Begonia sp.

Co lea sp.

Phyllarthron sp.

Phyllarthron sp.

Phyllarthron sp.

Canarium madagascariense Engl.

Canarium madagascariense Engl.

Canarium sp.

Protium sp.

Cinnamosma madagascariensis Danguy

Ascarina sp.

Calophyllum drouhardi H. Perrier

Calophyllum sp.

Mammea sp.

Mammea sp.

Mammea sp.

Mammea sp.

Mammea sp.

Ochrocarpos sp.

Ochrocarpos sp.

Ochrocarpos sp.

Ochrocarpos sp.

PJ 926

1875 75

PJ 934

1875 75

PJ 935

1875 75

NM 332

1625 75

NM 391

1875 75

NM 412

1875 75

NM 428

1875 75

PJ 765

775 75

PJ 865

1625 75

PJ 909

1875 75

PJ 922

1875 75

NM 346

1625 75

NM 374

1875 75

PJ 863

1625 75

PJ 923

1875 75

PJ 955

1875 75

NM 365

1875 75

NM 370

1875 75

PJ 819

1225 75

PJ 850

1625 75

PJ 870

1625 75

PJ 873

1625 75

PJ 932

1875 75

NM 256

775 75

NM 356

1625 75

NM 376

1875 75

NM 387

1875 75

NM 388

1875 75

NM 410

1875 75

PJ 774

775 75

PJ 821

1225 75

PJ 868

1625 75

PJ 896

1625 75

PJ 899

1625 75

PJ 921

1875 75

PJ 950

1875 75

NM 288

1225 75

NM 237

450 75

NM 317

1625 75

NM 345

1625 75

PJ 811

1225 75

NM 220

450 75

PJ 733

450 75

NM 294

1225 75

PJ 734

450 75

NM 247

775 75

NM 396

1875 75

PJ 841

1225 75

PJ 898

1625 75

NM 407

1875 75

NM 214

450 75

NM 323

1625 75

PJ 790

1225 75

PJ 797

1225 75

PJ 852

1625 75

PJ 904

1875 75

NM 242

450 75

NM 248

775 75

NM 250

775 75

NM 404

1875 75

98

FIELDIANA: ZOOLOGY

Appendix 3-4. Continued

Family

Genus and species

Collector*
and number

Altitude (m)t

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Clusiaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cunoniaceae

Cyperaceae

Cyperaceae

Droseraceae

Droseraceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Ebenaceae

Elaeocarpaceae

Elaeocarpaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Ericaceae

Eriocauleaceae

Erythroxylaceae

Erythroxylaceae

Erythroxylaceae

Erythroxylaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Ochrocarpos sp.

Psorospermum sp.

Psorospermum sp.

Psorospermum sp.

Symphonia microphylla (Cambess.) Vesque

Symphonia sp.

Symphonia sp.

Symphonia sp.

Symphonia sp.

Symphonia sp.

Weinmannia sp
Weinmannia sp
Weinmannia sp
Weinmannia sp
Weinmannia sp
Weinmannia sp
Weinmannia sp
Weinmannia sp
Cyperus sp.

Drosera sp.
Drosera sp.
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp
Diospyros sp

Elaeocarpus sp.

Sloanea rhodantha (Baker) Capuron

Agauria sp.

Agauria sp.

Wca sp.

Wca sp.

>/ca sp.

rica sp.

Wca sp.

Erica sp.

r/cfl sp.

>/ctf sp.

Vaccinium sp.

Vaccinium sp.

Vaccinium sp.

Vaccinium sp.

Eriocaulus sp.

Erythroxylum sp.

Erythroxylum sp.

Erythroxylum sp.

Erythroxylum sp.

Alchornea or Suregada sp.

Antidesma sp.

Antidesma sp.

PJ723

450 75

NM 277

1225 75

PJ 872

1625 75

PJ930

1875 75

PJ 890

1625 75

NM 216

450 75

NM 361

1875 75

PJ 727

450 75

PJ 801

1225 75

PJ 808

1225 75

NM 303

1225 75

NM 380

1875 75

NM 381

1875 75

NM 394

1875 75

NM 402

1875 75

NM423

1875 75

PJ 837

1225 75

PJ 897

1625 75

PJ959

1875 75

PJ908

1875 75

NM 369

1875 75

NM 363

1875 75

PJ916

1875 75

NM 268

1225 75

NM 287

1225 75

NM 302

1225 75

NM 316

1225 75

NM 354

1625 75

PJ 722

450 75

PJ 754

450 75

PJ 770

775 75

PJ 778

775 75

PJ782

1225 75

PJ 812

1225 75

PJ 826

1225 75

PJ 843

1225 75

NM 333

1625 75

PJ789

1225 75

NM 386

1875 75

PJ840

1225 75

NM 362

1875 75

NM 378

1875 75

NM 427

1875 75

PJ 799

1225 75

PJ903

1875 75

PJ958

1875 75

PJ967

1875 75

PJ969

1875 75

NM 340

1625 75

NM411

1875 75

NM 421

1875 75

PJ942

1875 75

NM 377

1875 75

NM 260

775 75

PJ 767

775 75

PJ 794

1225 75

PJ945

1875 75

PJ941

1875 75

PJ 721

450 75

PJ740

450 75

MESSMER ET AL.: VEGETATION

99

Appendix 3-4. Continued

Family

Genus and species

Collector*
and number

Altitude (m)t

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Euphorbiaceae

Fabaceae

Fabaceae

Flacourtiaceae

Flacourtiaceae

Flacourtiaceae

Flacourtiaceae

Gentianaceae

Gentianaceae

Gentianaceae

Gesneriaceae

Icacinaceae

Icacinaceae

Icacinaceae

Icacinaceae

Juncaceae

Juncaceae?

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Lauraceae

Loganiaceae

Loganiaceae

Loranthaceae

Loranthaceae

Antidesma sp.

Bridelia sp.

Croton sp.

Croton sp.

Croton sp.

Croton sp.

Croton sp.

Drypetes sp.

Drypetes sp.

Fluggea sp.

Macaranga cuspidata Baill.

Macaranga sp.

Macaranga sp.

Macaranga sp.

Macaranga sp.

Mallotus sp.

Phyllanthus sp.

Phyilanthus sp.

Phyllanthus sp.

Uapaca sp.

Uapaca sp.

Uapaca sp.

Uapaca sp.

Strongylodon sp.

Strongylodon sp.

Aphloia theiformis (Vahl) Benn.

Casearia sp.

Scolopia sp.

Scolopia sp.

Exacum sp.

Exacum sp.

Streptocarpus sp.
Cassinopsis sp.
Cassinopsis sp.

Appolonias sp.
Appolonias sp.
Cryptocarya sp.
Cryptocarya sp.
Cryptocarya sp.
Cryptocarya sp.
Cryptocarya sp.
Ocotea sp.
Ocotea sp.
Potameia sp.
Potameia sp.
Potameia sp.

Anthocleista madagascariensis Baker
Anthocleista madagascariensis Baker
Bakerella sp.
Bakerella sp.

PJ 741

450 75

PJ 736

450 75

NM 359

1875 75

NM 379

1875 75

PJ 793

1225 75

PJ 804

1225 75

PJ 953

1875 75

NM 270

1225 75

NM 273

1225 75

PJ 717

450 75

NM 234

450 75

NM 213

450 75

NM 221

450 75

NM 292

1225 75

PJ 885

1625 75

PJ 730

450 75

NM 239

450 75

NM 298

1225 75

PJ 779

775 75

NM 254

775 75

NM 279

1225 75

NM 313

1225 75

NM 330

1625 75

PJ 743

450 75

NM 264

775 75

PJ 751

450 75

PJ 857

1625 75

PJ 759

775 75

PJ 894

1625 75

PJ 928

1875 75

NM 274

1225 75

NM 390

1875 75

PJ 920

1875 75

PJ 745

450 75

NM 358

1875 75

PJ 817

1225 75

PJ 829

1225 75

PJ 844

1225 75

PJ 907

1875 75

NM 401

1875 75

NM 397

1875 75

PJ 951

1875 75

NM 276

1225 75

NM 319

1625 75

PJ 758

775 75

PJ 785

1225 75

PJ 882

1625 75

NM 236

450 75

NM 311

1225 75

NM 284

1225 75

NM 326

1625 75

PJ 946

1875 75

NM 285

1225 75

PJ 825

1225 75

PJ 845

1225 75

PJ 867

1625 75

PJ 851

1625 75

PJ 948

1875 75

NM 297

1225 75

NM 395

1875 75

100

FIELDIANA: ZOOLOGY

Appendix 3-4. Continued

Family

Genus and species

Collector*
and number

Altitude (m)t

Loranthaceae

Loranthaceae

Loranthaceae

Melanophyllaceae

Melastomataceae

Melastomataceae

Melastomataceae

Melastomataceae

Melastomataceae

Melastomataceae

Melastomataceae

Melastomataceae

Meliaceae

Meliaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Monimiaceae

Moraceae

Moraceae

Moraceae

Moraceae

Moraceae

Moraceae

Moraceae

Moraceae

Moraceae

Myricaceae

Myricaceae

Myricaceae

Myricaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

Myrsinaceae

DC.

Bakerella sp.

Melanophylla humbertiana Keraudren
Dichaetanthera sp.
Dichaetanthera sp.
Dichaetanthera sp.
Dionycha sp.
Memecylon sp.
Memecylon sp.

Malleastrum sp.

Malleastrum sp.

Decarydendron sp.

Ephippiandra madagascariensis (Cavaco)

Ephippiandra perrieri (Cavaco) Lorence

Tambourissa cf. parvifolia Baker

Tambourissa cf. purpurea (Tul.) A. 1

Tambourissa purpurea (Tul.) A. DC

Tambourissa sp.

Tambourissa sp.

Tambourissa sp.

Tambourissa sp.

Tambourissa sp.

Tambourissa thouvenotii Danguy

Dorstenia sp.

Fie us politoria Lam.

Ficus politoria Lam.

Ficus sp.

Ficus sp.

Ficus sp.

Ficus sp.

Ficus tiliaefolia Baker

Trilepisium madagascariense DC.

Myrica cf. spathulata Mirb.

Myrica phillyreaefolia Baker

Myrica phillyreaefolia Baker

Myrica sp.

Embelia sp.

Embelia sp.

Maesa sp.

Monoporus sp.

Monoporus sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Oncostemum sp.

Lorence

NM 406

1875 75

PJ 863

1625 75

PJ956

1875 75

PJ 732

450 75

NM 251

775 75

NM 336

1625 75

PJ 773

775 75

PJ940

1875 75

NM 241

450 75

PJ 744

450 75

NM 348

1625 75

NM 416

1875 75

NM 212

450 75

PJ 756

775 75

PJ 839

1225 75

PJ 818

1225 75

PJ 905

1875 75

PJ 884

1625 75

NM 249

775 75

NM 307

1225 75

NM 301

1225 75

PJ 731

450 75

PJ 772

775 75

PJ 788

1225 75

PJ 810

1225 75

PJ 747

450 75

PJ 775

775 75

NM 261

775 75

PJ 777

775 75

NM 226

450 75

NM 230

450 75

NM 233

450 75

PJ 724

450 75

PJ 755

450 75

NM 222

450 75

PJ 876

1625 75

PJ 802

1225 75

PJ 875

1625 75

PJ929

1875 75

PJ 834

1225 75

PJ939

1875 75

PJ 816

1225 75

PJ 856

1625 75

PJ947

1875 75

NM 224

450 75

NM 244

775 75

NM 339

1625 75

NM 393

1875 75

NM 403

1875 75

NM 409

1875 75

NM 415

1875 75

NM420

1875 75

NM 425

1875 75

PJ 718

450 75

PJ 760

775 75

PJ 780

775 75

PJ 792

1225 75

PJ 803

1225 75

PJ 832

1225 75

PJ 891

1625 75

MESSMER ET AL.: VEGETATION

101

Appendix 3-4. Continued

Collector*

Family

Genus and species

and number

Altitude (m)t

Myrsinaceae

Oncostemum sp.

PJ937

1875

75

Myrsinaceae

PJ 860

1625

75

Myrsinaceae

PJ957

1875

75

Myrtaceae

Eugenia sp.

PJ918

1875

75

Myrtaceae

Syzygium sp.

NM 282

1225

75

Myrtaceae

Syzygium sp.

NM 304

1225

75

Myrtaceae

Syzygium sp.

NM 329

1625

75

Myrtaceae

Syzygium sp.

NM424

1875

75

Myrtaceae

Syzygium sp.

PJ 858

1625

75

Myrtaceae

Syzygium sp.

PJ933

1875

75

Myrtaceae

Syzygium sp.

PJ960

1875

75

Myrtaceae

Syzygium sp.

PJ961

1875

75

Myrtaceae

Syzygium sp.

PJ962

1875

75

Myrtaceae

Syzygium sp.

PJ963

1875

75

Oleaceae

Noronhia sp.

PJ 720

450

75

Oleaceae

Noronhia sp.

PJ 729

450

75

Orchidaceae

Angraecum sp.

NM 228

450

75

Orchidaceae

Angraecum sp.

NM 229

450

75

Orchidaceae

Angraecum sp.

NM 255

775

75

Orchidaceae

Angraecum sp.

PJ 889

1625

75

Orchidaceae

Bulbophyllum sp.

PJ 831

1225

75

Orchidaceae

Bulbophyllum sp.

PJ906

1875

75

Orchidaceae

Calanthes sp.

PJ 886

1625

75

Orchidaceae

Cynorchis sp.

NM 266

775

75

Orchidaceae

Cynorchis sp.

NM 372

1875

75

Orchidaceae

NM 352

1625

75

Orchidaceae

NM 364

1875

75

Orchidaceae

PJ 887

1625

75

Orchidaceae

PJ910

1875

75

Piperaceae

Peperomia sp.

NM 210

450

75

Piperaceae

Peperomia sp.

NM 218

450

75

Piperaceae

Peperomia sp.

NM 219

450

75

Piperaceae

Peperomia sp.

NM 243

450

75

Piperaceae

Peperomia sp.

NM 259

775

75

Piperaceae

Peperomia sp.

NM 289

1225

75

Piperaceae

Peperomia sp.

NM 295

1225

75

Piperaceae

Peperomia sp.

NM 308

1225

75

Piperaceae

Peperomia sp.

NM 309

1225

75

Piperaceae

Peperomia sp.

NM 353

1625

75

Piperaceae

Peperomia sp.

NM 360

1875

75

Piperaceae

Pzp^r sp.

NM 231

450

75

Piperaceae

P//?er sp.

NM 240

450

75

Piperaceae

Pz/jer sp.

NM 296

1225

75

Pittosporaceae

Pittosporum ambrense Cufod.

PJ 815

1225

75

Pittosporaceae

Pittosporum ambrense Cufod.

PJ 833

1225

75

Pittosporaceae

Pittosporum cf. verticillatum Bojer

NM 291

1225

75

Pittosporaceae

Pittosporum humbertii Cufod.

PJ968

1875

75

Pittosporaceae

Pittosporum polysperrnum Tul.

PJ 719

450

75

Pittosporaceae

Pittosporum polysperrnum Tul.

PJ 781

775

75

Pittosporaceae

Pittosporum sp.

NM 217

450

75

Pittosporaceae

Pittosporum verticillatum Bojer

PJ 880

1625

75

Poaceae

NM 280

1225

75

Poaceae

PJ 749

450

75

Poaceae

PJ 912

1875

75

Poaceae

PJ913

1875

75

Poaceae

PJ 915

1875

75

Proteaceae

Faurea forficuliflora Baker var. elliptica Humbert

PJ 864

1625

75

Rhizophoraceae

Cassipourea sp.

NM 278

1225

75

Rhizophoraceae

Cassipourea sp.

NM 322

1625

75

Rhizophoraceae

Cassipourea sp.

PJ 828

1225

75

102

FIELDIANA: ZOOLOGY

Appendix 3-4. Continued

Collector*

Family

Genus and species

and number

Altitude (m)t

Rhizophoraceae

Cassipourea sp.

PJ 881

1625

75

Rhizophoraceae

Macarisia sp.

PJ 854

1625

75

Rhizophoraceae

PJ 805

1225

75

Rubiaceae

Alberta minor Baill.

NM 318

1625

75

Rubiaceae

Alberta minor Baill.

NM 331

1625

75

Rubiaceae

Alberta minor Baill.

NM426

1875

75

Rubiaceae

Alberta sambiranensis Cavaco

NM 293

1225

75

Rubiaceae

Canthium sp.

NM 246

775

75

Rubiaceae

Canthium sp.

NM 269

1225

75

Rubiaceae

Canthium sp.

NM 290

1225

75

Rubiaceae

Canthium sp.

PJ 764

775

75

Rubiaceae

Canthium sp.

PJ 807

1225

75

Rubiaceae

Canthium sp.

PJ 842

1225

75

Rubiaceae

Canthium sp.

PJ 892

1625

75

Rubiaceae

cf. Chapelieria sp.

NM 347

1625

75

Rubiaceae

cf. Chapelieria sp.

PJ 813

1225

75

Rubiaceae

cf. Chapelieria sp.

PJ 846

1625

75

Rubiaceae

cf. Chapelieria sp.

PJ 847

1625

75

Rubiaceae

Danais cf. tsaratananensis Homolle

NM 341

1625

75

Rubiaceae

Gaertnera macrostipula Baker

PJ 796

1225

75

Rubiaceae

Gaertnera sp.

NM 245

775

75

Rubiaceae

Gaertnera sp.

NM 338

1625

75

Rubiaceae

Gaertnera sp.

NM405

1875

75

Rubiaceae

Gaertnera sp.

PJ742

450

75

Rubiaceae

Gaertnera sp.

PJ 809

1225

75

Rubiaceae

Gaertnera sp.

PJ 848

1625

75

Rubiaceae

Gaertnera uniflora ined.

NM 300

1225

75

Rubiaceae

Hyperacanthus sp.

PJ 762

775

75

Rubiaceae

Ixora sp.

NM 252

775

75

Rubiaceae

Ixora sp.

NM 267

1225

75

Rubiaceae

Ixora sp.

NM 349

1625

75

Rubiaceae

Ixora sp.

PJ 735

450

75

Rubiaceae

Ixora sp.

PJ 883

1625

75

Rubiaceae

Lemyrea sp.

PJ 784

1225

75

Rubiaceae

Mapouria pyrrotricha Bremek.

NM 299

1225

75

Rubiaceae

Morinda sp.

PJ 769

775

75

Rubiaceae

Mussaenda erectiloba Wernham

NM 325

1625

75

Rubiaceae

Nematostylis anthophylla (A. Rich.) Baill.

NM 367

1875

75

Rubiaceae

Nematostylis anthophylla (A. Rich.) Baill.

NM 399

1875

75

Rubiaceae

Pauridiantha paucinervis

NM 225

450

75

Rubiaceae

Payera beondrokensis (Humbert) Buchner & Puff

PJ900

1875

75

Rubiaceae

Payera sp.

PJ 822

1225

75

Rubiaceae

Psychotria cf. ankafinensis (K. Schum.) Bremek.

NM400

1875

75

Rubiaceae

Psychotria lokohensis Bremek.

PJ716

450

75

Rubiaceae

Psychotria sp.

NM272

1225

75

Rubiaceae

Psychotria sp.

NM 355

1625

75

Rubiaceae

Psychotria sp.

NM 417

1875

75

Rubiaceae

Psychotria sp.

NM418

1875

75

Rubiaceae

Psychotria sp.

PJ715

450

75

Rubiaceae

Psychotria sp.

PJ738

450

75

Rubiaceae

Psychotria sp.

PJ 814

1225

75

Rubiaceae

Psychotria sp.

PJ 824

1225

75

Rubiaceae

Psychotria sp.

PJ 949

1875

75

Rubiaceae

Sabicea diversifolia Pers.

NM 258

775

75

Rubiaceae

Saldinia sp.

PJ739

450

75

Rubiaceae

Schismatoclada cf. farahimpensis Homolle

PJ 737

450

75

Rubiaceae

Schismatoclada sp.

NM 342

1625

75

Rubiaceae

Schismatoclada sp.

NM 392

1875

75

Rubiaceae

Schismatoclada sp.

PJ 965

1875

75

Rubiaceae

Schismatoclada sp.

PJ966

1875

75

MESSMER ET AL.: VEGETATION

103

Appendix 3-4. Continued

Familv

Genus and species

Collector*
and number

Altitude (m)i

Rubiaceae

Rubiaceae

Rubiaceae

Rubiaceae

Rubiaceae

Rubiaceae

Rutaceae

Rutaceae

Rutaceae

Rutaceae

Rutaceae

Rutaceae

Sapindaceae

Sapindaceae

Sapindaceae

Sapindaceae

Sapindaceae

Sapindaceae

Sapindaceae

Sapotaceae

Sapotaceae

Sapotaceae

Sapotaceae

Sapotaceae

Scrophulariaceae

Sterculiaceae

Sterculiaceae

Sterculiaceae

Sterculiaceae

Thymeleaceae

Thymeleaceae

Thymeleaceae

Tiliaceae

Ulmaceae

Urticaceae

Urticaceae

Urticaceae?

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Verbenaceae

Viscaceae

Xyridaceae

Xyridaceae

Xyridaceae

Xyridaceae

Tarenna sp.
Tare una sp.

Citrus sp.

Evodia or Ivodea sp.
Evodia or Ivodea sp.
Evodia sp.
Vepris sp.

Tina sp.

Tina sp.

Tina sp.

Tina sp.

Tina sp.

Tina sp.

Tina striata

Faucherea sp.

Faucherea sp.

Faucherea sp.

Faucherea thouvenotii Lecomte

Faucherea thouvenotii Lecomte

Dombeya sp.

Dombeya sp.

Dombeya sp.

Dombeya sp.

Atemnosiphon coriaceus Leandri

Atemnosiphon coriaceus Leandri

Octolepis sp.

Greuia sp.

Clerodendron sp.
Clerodendron sp.
Clerodendron sp.
Clerodendron sp.
Clerodendron sp.

Viscum sp.
Xyris sp.
Xvm sp.
Xyris sp.
Xvm sp.

PJ 766

775 75

PJ 835

1225 75

NM 324

1625 75

PJ 931

1875 75

PJ 943

1875 75

PJ 964

1875 75

NM 232

450 75

PJ 877

1625 75

PJ 878

1625 75

NM 398

1875 75

NM 286

1225 75

PJ 879

1625 75

NM 275

1225 75

NM 315

1225 75

NM 320

1625 75

NM 327

1625 75

PJ 791

1225 75

PJ 823

1225 75

PJ 786

1225 75

NM 253

775 75

NM 271

1225 75

NM 335

1625 75

PJ 838

1225 75

NM 283

1225 75

NM 351

1625 75

NM 373

1875 75

NM419

1875 75

PJ 763

775 75

PJ 893

1625 75

NM 321

1625 75

PJ 795

1225 75

PJ 853

1625 75

PJ 753

450 75

PJ 750

450 75

NM 265

775 75

PJ 888

1625 75

NM 262

775 75

NM 389

1875 75

PJ 783

1225 75

PJ 901

1875 75

PJ927

1875 75

PJ 954

1875 75

NM 328

1625 75

NM 357

1875 75

PJ 830

1225 75

PJ 861

1625 75

PJ 771

775 75

NM 366

1875 75

NM 371

1875 75

PJ 869

1625 75

PJ917

1875 75

NM 368

1875 75

PJ866

1625 75

PJ 911

1875 75

PJ936

1875 75

PJ952

1875 75

Collectors: NM. Nathalie Messmer: PJ. Pierre Jules Rakotomalaza.
The altitudes refer to the five elevational zones.

104

FIELDIANA: ZOOLOGY

Chapter 4

The First Beckeriella Williston from the Afrotropical
Region: Two New Species from Madagascar
(Diptera: Ephydridae)

Wayne N. Mathis 1 and David A. Grimaldi 2

Abstract

Two new species of the shore-fly genus Beckeriella are described from specimens collected
on the island of Madagascar. These are the first species of Beckeriella from the Afrotropical
Region, and aside from B. filipina (from the Philippines), they are the first species of the genus
to be described from regions outside of the New World tropics, where the genus has its greatest
diversity.

Resume

Deux nouvelles especes du Madagascar du genre "la mouche de la plage" Beckeriella sont
decrites. Ce sont les premieres especes du genre Beckeriella recoltees dans la region afrotrop-
icale. Hors B. filipina (des Phillipines) les especes decrites ici sont les premieres qui proviennent
des regions autres que les neotropiques, ou ce genre a actuellement la plus grande diversite.

Introduction

Beckeriella Williston is an uncommonly col-
lected genus of shore flies that, until recently, was
represented by 10 New World species (Mathis &
Zatwarnicki, 1995). Four years ago, however, Li-
zarralde de Grosso (1994) described B. filipina
from the Philippines, which substantively and
somewhat anomalously increased the known dis-
tribution for the genus. Such an apparent disjunct
distribution between the Philippines and the trop-
ical neotropics was unknown among the Ephydri-
dae. Here we extend even further the known
worldwide distribution and describe two new spe-
cies from Madagascar, the first of this genus from

1 Department of Entomology, NHB 169, Smithsonian
Institution, Washington, D.C. 20560, U.S.A.

2 Division of Invertebrate Zoology, American Muse-
um of Natural History, Central Park West at 79th St.,
New York, NY 10024-5192, U.S.A.

the Afrotropical Region. This further expands the
known distribution of Beckeriella and alters our
ideas about the historical origins and phylogeny
of the genus.

Methods

The terminology and methods used in this
study were explained previously (Mathis, 1990).
Because of the small size of the specimens, study
and illustration of the male terminalia required the
use of a compound microscope. To ensure effec-
tive communication about structures of the male
terminalia, we have adopted the terminology of
other workers in Ephydridae (see references in
Mathis, 1986). Usage of these terms, however,
should not be taken as an endorsement of them
from a theoretical or morphological view over al-
ternatives that have been proposed (Cumming et

MATHIS & GRIMALDI: SHORE FLIES BECKERIELLA

105

al., 1995; Griffiths, 1972; McAlpine, 1981). Rath-
er, we are deferring to tradition until the morpho-
logical issues are better resolved.

Three ratios (one cephalic, two venational) are
commonly used in the descriptions and are de-
fined here for the convenience of the user (ratios
are ranges based on three specimens): 1. Gena-to-
eye ratio is the genal height measured at the max-
imum eye height divided by the eye height. 2.
Costal vein ratio is the straight-line distance be-
tween the apices of veins R 2+3 and R 4 ^ s /distance
between the apices of veins R, and R 2 ^. 3. M
vein ratio is the straight-line distance along M be-
tween crossveins dm-cu and r-m/distance apicad
of crossvein dm-cu.

Tribe Gastropini Cresson

Gastropini Cresson, 1949:250 (as Gastropsini).

Type Genus Gastrops Williston, 1897 (Za-
twarnicki, 1995:127-130 [world catalog]).

Discussion Williston (1897) described both
genera, Beckeriella and Gastrops, that now make
up the tribe Gastropini, which until recently was
placed in the subfamily Parydrinae (Cresson,
1949; Lizarralde de Grosso, 1989; Wirth, 1968).
Currently the tribe is placed in the subfamily
Gymnomyzinae, based on Zatwarnicki's (1992)
assessment of characters of the male terminalia.
Although the tribe is undoubtedly monophyletic
(Mathis, 1977), other aspects concerning the phy-
logenetic relationships of the tribe, such as iden-
tification of the tribe's sister group and the mono-
phyly of the two included genera, are unresolved.
Two colleagues, Stark (pers. comm.) and Lizar-
ralde de Grosso (pers. comm.), are now studying
these phylogenetic issues as part of more compre-
hensive research on the tribe. Thus, we defer com-
ment on the phylogeny and taxonomic status of
the included genera and recognize Beckeriella as
described previously and diagnosed herein.

Genus Beckeriella Williston

Beckeriella Williston, 1897:2.

Type Species Ephydra bispinosa Thomson,
1868 (=Scatophaga bispinosa Fabricius, 1805),
monotypy. (Cresson, 1934:201-205 [review];
Wirth, 1968:20 [Neotropical catalog]; Lizarralde
de Grosso, 1986:133-137 [species description];

1990:143-146 [species description]; 1991a:39-42
[species description]; 1991b:45-48 [species de-
scription]; 1992:229-233 [species description];
1994:57-59 [species description, Philippines];
Mathis & Zatwarnicki, 1995:27-129 [world cat-
alog]).

Diagnosis A tribe of the subfamily Gymno-
myzinae that is distinguished from other tribes by
the following combination of characters: moder-
ately small to large shore flies, length 2.5-5.5
mm.

Head Frons rectangular, wider than long, ver-
tex at level beyond posterior margin of posterior
ocelli; pseudopostocellar setae greatly reduced or
lacking; ocellar seta well developed, subequal to
outer vertical seta, proclinate, slightly divergent;
fronto-orbital seta 2, both proclinate, only anterior
setae usually well developed, oriented anterolat-
erally, posterior seta much smaller or lacking; out-
er vertical seta well developed; inner vertical seta
weakly developed to lacking, if present much
smaller than outer seta and with medioproclinate
orientation. Scape and pedicel comparatively long
and semiporrect; flagellomere 1, comparatively
long, greatly elongate and pendulous in some spe-
cies, apex rounded; pedicle lacking a well-devel-
oped dorsoapical seta; flagellum long, bearing 6-
12 dorsal rays along length. Face in profile usu-
ally concave, sometimes with dorsal half shallow-
ly tumescent, usually with a transverse crease near
midheight, usually bare of microtomentum, shiny,
bearing a single seta and smaller setulae laterally
near margin with parafacial, rarely bearing a 2nd
seta ventrad from larger seta. Eye somewhat pro-
jected laterally, bulbous, horizontally ovate
(bluntly rounded anteriorly in lateral view, more
narrowly rounded posterodorsally) or vertically
ovate. Anterior oral margin more or less straight;
clypeus wide, bandlike, short, partially to mostly
concealed by ventral margin of face and gena; ge-
nal seta 1, frequently reduced in size.

Thorax Generally dark-colored; scutellum
usually moderately flat, rectangular to trapezoidal,
only apical scutellar seta well developed, these
sometimes arising from small tubercles; acrosti-
chal setulae minute, usually in 2 rows; only the
posteriormost dorsocentral seta well developed,
slightly displaced laterally; no presutural or post-
sutural supra-alar seta; postalar seta 1; proprono-
tum setulose but lacking a large seta; notoplueron
usually with 2 setae near ventral margin, posterior
seta stronger, anterior seta sometimes lacking;
anepisternum bearing 1 large seta along posterior
margin; katepisternum bearing 1 large seta and

106

FIELDIANA: ZOOLOGY

with anterior 2 h usually bare of microtomentum,
similar to anteroventral corner of anepisternum,
shiny; otherwise pleuron thinly microtomentose.
Wing usually infuscate, sometimes very dark, to
maculate; vein R 2 . 3 curved anteriorly at apex rath-
er abruptly, often bearing a subapical stump vein;
frequently with some veins sinuous, especially
veins R 2 + 3 , R 4+ 5, and M. Halter with knob blackish
brown. Femora usually brown to black; fore- and
especially hind femora sometimes swollen; fore-
femur bearing numerous long setae along poster-
o ventral surface; midfemur bearing numerous
long setae along anteroventral surface; basal tar-
someres yellow, apical 2-3 brownish black.

Abdomen Strongly convex, domelike, thinly
to moderately invested with microtomentum,
sometimes more so toward posterior margin, to
nearly bare, subshiny to shiny, often with some
metallic luster; tergites with lateroventral margins
sharply defined, often with cuticular microsculp-
turing, surface appearing shallowly wrinkled or
puckered; 2nd tergite with patch of dorsoerect se-
tulae laterally.

Distribution With the exception of the two
species described here and B. filipina (from the
Philippines), Beckeriella is known only from the
New World tropics, where the genus currently has
its greatest diversity, with 10 species. Many un-
described species are already known and available
in collections (Stark, pers. comm.), and undoubt-
edly numerous others remain to be discovered.
Although the shore flies of the Afrotropical Re-
gion have not been well collected, we find it
somewhat anomalous that the first species of
Beckeriella were found on Madagascar and not in
continental Africa. On Madagascar, the two spe-
cies were collected at higher elevation, above
1000 m, and from the same province (Antsiran-
ana).

Discussion Beckeriella is apparently of an-
cient origin and is the only identified genus of
shore flies that is represented by fossils in amber
(Evenhuis, 1994). The fossils, which are from Do-
minican amber of unknown provenance, represent
a few species and date from the Oligocene/Mio-
cene epochs. Stark (pers. comm.) is now studying
and describing these species.

Beckeriella fasciata, new species

Figures 4-1, 4-2a,b

Description A large shore fly (Fig. 4-1),
length 5.2 mm; head generally yellow; thorax and

abdomen black, subshiny to shiny; wing generally
dark brown with subapical transverse white band.

Head (Fig. 4-2a) Generally yellow; meso-
frons rectangular, longer than wide, brown; fron-
to-orbits brown. Inner vertical seta greatly re-
duced or lacking; fronto-orbital setae 2, posterior
seta short, much less than half length of anterior
seta, anterior seta slightly longer than outer ver-
tical seta. Scape yellowish brown basally becom-
ing brown apically, length distinctly shorter than
pedicle; pedicle brown with some gray microto-
mentum dorsally; flagellomere 1 dark brown on
dorsal half, yellow on ventral half both laterally
and medially, length about 2.5 X height; flagel-
lomere bearing 1 1 dorsal rays. Face concave in
lateral view, yellow except for wide stripe just
ventrad of antennal bases and a thinner stripe at
midheight of facial concavity. Eye transversely
ovate, anterior margin bluntly rounded, tapered
posteriorly to more narrowed posterior margin.
Gena yellow, moderately high, gena-to-eye ratio
0.39. Clypeus yellow, wide, short, largely con-
cealed in oral cavity. Mouthparts yellow.

Thorax Generally black; scutum subshiny,
thinly invested with brown microtomentum anter-
omedially, anterior and anterolateral margins of
scutum lightly gray microtomentose, microtomen-
tum becoming sparser and appearing shinier pos-
teriorly until just before posterior margin, which
is narrowly but more densely microtomentose,
brown; scutellum very sparsely and uniformly
brown microtomentose, rectangular (Fig. 4-2b),
wider than long, with posterior margin broadly
truncate, apical scutellar setae arising from short
tubercles at posterolateral corners of scutellum,
about as long as length of scutellum; pleuron
black, mostly uniformly sparsely microtomentose
except for shiny, bare anteroventral corner of ane-
pisternum and anterior 2 h of katepisternum. Wing
mostly brown, slightly darker basally, with irreg-
ular (basal and apical margins) subapical trans-
verse white hand at level of apex of vein R 2+3 and
continued posteriorly just apical of crossvein dm-
cu; crossvein r-m slightly whitened medially; vein
R 2+3 sinuous, curved sharply toward costa at apex,
short, making costal section III as long as section
II; vein R 4+5 sinuous; crossvein dm-cu longer than
apex of vein CuA, apicad of crossvein dm-cu;
costal vein ratio 0.87; M vein ratio 0.60; alula
short and wide, fringing setulae as long as alular
height; knob of halter blackish brown. Legs most-
ly brownish black to black, sometimes with shiny
surfaces, only foretibia and basal 3 tarsomeres

MATHIS & GRIMALDI: SHORE FLIES BECKERIELLA

107

Fig. 4-1. Beckeriella fasciata. Habitus, lateral view.

yellow; apices of mid- and hind tibiae brown,
slightly lighter in color than femora.

Abdomen Appearing domelike, generally
black, subshiny, thinly microtome ntose; 2nd ter-
gite with lateral patches of dorsoerect setae; lat-
eral margins of tergites sharply defined.

Type Material The holotype female is la-
beled "Madagascar-Est dct. Sambava R.N.
[now Pare National de Marojejy] XII Marojejy-
Ouest 1140 m XI-59 [Nov 1959] P.Soga/INSTI-
TUT SCIENTIFIQUE MADAGASCAR [blue;
black margin]." The holotype is double-mount-
ed (minuten in rectangular block or pith), is in
good condition (right flagellomere 1 missing,
left wing torn apically, some tarsi missing, an-

tennae lost soon after illustrating), and is de-
posited in the Natal Museum (Pietermartzburg,
South Africa).

Distribution Afrotropical: Madagascar (An-
tsiranana).

Etymology The specific epithet, fasciata, re-
fers to the subapical white fascia or band on an
otherwise brown to dark brown wing.

Diagnosis This species is distinguished from
congeners, especially B. maculata, by its large
size (5.2 mm), the largely brown wing with a sub-
apical white transverse band, and its body color-
ation (see description).

Remarks This species is represented by a sin-
gle female.

108

FIELDIANA: ZOOLOGY

Fig. 4-2. a, b: Beckeriella fasciata. a, Head, anterior view, b, Scutellum, dorsal view, c-e: Beckeriella maculata.
c, Head, anterior view, d, Scutellum, dorsal view, e, Male genitalia, lateral view.

Beckeriella maculata, new species

Figures 4-2c-e, 4-3 to 4-5

Description Moderately small shore fly,
length 2.5-2.70 mm; head generally yellow; tho-
rax and abdomen black, subshiny to shiny; wing
generally dark brown on basal half, apical half
white with maculate pattern.

Head (Fig. 4-2c) Generally yellow; meso-
frons rectangular, slightly wider than long; fronto-
orbits yellow. Inner vertical seta well developed,
equal in length to fronto-orbital seta; fronto-orbit-
al setae 2, posterior seta very short, much less
than half length of anterior seta, anterior seta
slightly longer than outer vertical seta. Scape and
pedicle yellow with some grayish microtomentum
dorsally; length of scape distinctly shorter than

pedicle; flagellomere 1 mostly yellow, lateral sur-
face with brown stripe extended from anterodorsal
corner to base, yellow on dorsal and ventral sides
of stripe, median surface with only anterodorsal
corner brown, otherwise yellow, length about
twice height; flagellomere bearing 9-10 dorsal
rays. Face shiny, bare of microtomentum, concave
in lateral view with distinct transverse crease at
midheight, with a transverse brown stripe at
crease, otherwise whitish yellow. Eye transversely
ovate, anterior margin bluntly rounded, tapered
posteriorly to more narrowed posterior margin.
Gena yellow, moderately low, gena-to-eye ratio
0.26-0.30. Clypeus yellow, wide, short, largely
concealed in oral cavity. Mouthparts yellow.

Thorax Generally black; scutum subshiny,
anterior and anterolateral margins of scutum light-

MATHIS & GRIMALDI: SHORE FLIES BECKERIELLA

109

Fig. 4-3. Beckeriella maculata. Habitus, lateral view.

ly gray microtomentose, posterior half to % of
scutum mostly with brown microtomentum, with
long gray linear spot on median side of dorsocen-
tral track that extends from anterior margin pos-
teriorly for % length of scutum and 2 linear spots
laterally on either side of transverse suture; scu-
tellum with dorsal surface finely granulose, most-
ly black with some bluish black reflections, very
sparsely brown microtomentose, trapezoidal (Fig.
4-2d), about as long as wide, gradually tapered
posteriorly, with posterior margin emarginate me-

dially, apical scutellar setae arising from postero-
lateral corners, without tubercles at base, subapi-
cal scutellar setae about % length of apical setae,
inserted laterally at apical %. Wing mostly brown
on basal half, dark brown basally, becoming
slightly and gradually lighter toward apical half;
apical half of wing maculate, mostly hyaline with
dark brown irregular spots and zigzag and C-
shaped bands; vein R 2 . 3 sinuous, curved sharply
toward costa at apex, bearing a subapical stump
vein that is oriented posteriorly, vein R 2 . 3 short,

110

FIELDIANA: ZOOLOGY

Fig. 4-4. Beckeriella maculata. Male terminalia,
ventral view, including epandrium, cerci, and sternites
2-5 (5th divided).

making costal section III as long as section II;
vein R 4+5 sinuous; cross vein dm-cu longer than
apex of vein CuA, apicad of crossvein dm-cu;
costal vein ratio 1.15-1.18; M vein ratio 0.55-
0.57; alula short and wide, fringing setulae black,
longer than alular height; knob of halter blackish
brown. Legs with coxae, trochanters, and femora
black; tibiae mostly yellow, with dorsal surfaces
of mid- and hind tibiae partially dark brown; basal
3 tarsomeres yellow, apical 2 blackish brown; api-

ces of mid- and hind tibiae brown, slightly lighter
in color than femora.

Abdomen Generally black, subshiny to
shiny, with some dark blue reflections; generally
sparsely gray microtomentose, especially on an-
terior % of tergites, microtomentum becoming
denser submarginally along posterior margin of
tergites. Male abdomen (Fig. 4-4): sternites 2-4
rounded, bearing setulae along lateral margins;
sternite 2 with small anteromedial papilla; 5th
sternite subdivided; epandrium in ventral view
widest at midheight, angulate laterally at greatest
width; cerci in ventral view subelliptical, medial
margin sinuous, wider anteriorly; aedeagus (Fig.
4-2e) tubular; aedeagal apodeme (Fig. 4-2e) with
long, narrow keel; granite (Fig. 4-2e) with apex
recurved, pointed; hypandrium a somewhat rect-
angular sclerite with anterior margin shallowly
rounded. Female abdomen (Fig. 4-5): Tergites
generally wide, 2-4 comparatively long, tergites
5-8 short, 4th with 2 narrow patches of microto-
mentum toward posterior margin on either side of
median; sternites 2-4 subquadrate, angles round-
ed; sternites 5-7 conspicuously wider than long,
width of 7th about Vi 5th, 8th sternite subdivided,
each sclerite circular; female ventral receptacle
(Fig. 4-5) with operculum bell-shaped, higher
than wide, extended process J-shaped, as long as
height of operculum.

Type Material The holotype male is labeled
"MADAGASCAR: Antsiranana, Marojejy, 1 1 km
NW Manantenina, X.25-XI.3.1996 1,225 m, E.
Quinter." The holotype is double mounted (glued
to a paper triangle), is in good condition, and is
deposited in the American Museum of Natural
History (AMNH). Thirteen paratypes bear the
same locality label data as the holotype (9
AMNH, 4 United States National Museum
[USNM]). All specimens of the type series were
collected in pan traps set at the side of a stream.

Distribution Afrotropical: Madagascar (An-
tsiranana).

Etymology The specific epithet, maculata,
refers to the dappled or maculate apical half of
the wing.

Key to Malagasy Species of Beckeriella

1. Wing mostly brown with irregular subapical
transverse white band; scutellum rectangular,
wider than long, posterior margin broadly trun-
cate; inner vertical seta greatly reduced, length

MATHIS & GRIMALDI: SHORE FLIES BECKERIELLA

111

Fig. 4-5. Beckeriella maculata. Female terminalia. posterior view, showing apical tergites and sternites. and
ventral receptacle, lateral view.

much less than half that of outer vertical seta

B. fasciata, new species

Wing with basal half brown and apical half
maculate, mostly hyaline with dark brown
spots; scutellum trapezoidal, about as long as
wide, tapered posteriorly, posterior margin
emarginate medially; inner vertical seta well
developed, equal in length to fronto-orbital
seta B. maculata, new species

AMNH. The holotype of B. fasciata was loaned
to us by the Natal Museum, Pietermartzburg,
South Africa (David A. Barraclough). Sally
Goodman made the habitus illustrations and
helped with line drawings. For critically review-
ing a draft of this paper, we thank Volker Holl-
mann-Schirrmacher and Julian Stark.

Acknowledgments

We are grateful to Eric Quinter. collector of the
specimens of B. maculata, for making the type
series of this species available to us. Funding for
Eric's expeditionary work on Madagascar was
provided by World Wide Fund for Nature (WWF)
and the Center for Biodiversity and Conservation.

Literature Cited

Cresson. B. T. Jr. 1934. Descriptions of new genera
and species of the dipterous family Ephydridae. XI.
Transactions of the American Entomological Society,
60: 199-222.

1949. A systematic annotated arrangement of

the genera and species of the North American Ephy-
dridae (Diptera). IV. The Subfamily Napaeinae. Trans-
actions of the American Entomological Societv. 74:
225-260.
Claiming. J. M.. B. J. Sinclair, and D. M. Wood. 1995.

112

FTELDIANA: ZOOLOGY

Homology and phylogenetic implications of male gen-
italia in Diptera Eremoneura. Entomologica Scan-
dinavica, 26: 120-151.
Evenhuis, N. L. 1994. Catalogue of the Fossil Flies of
the World (Insecta: Diptera). Backhuys Publishers,
Leiden, 600 pp.

Griffiths, G. C. D. 1972. The phylogenetic classifica-
tion of Diptera Cyclorrhapha with special reference to
the structure of the male postabdomen. Series Ento-
mologica, 8: 1-340, W. Junk, The Hague.

Lizarralde de Grosso, M. S. 1986. Dos nuevas espe-
cies de Beckeriella Williston (Diptera Ephydridae).
Acta Zoologica Lilloana, 38(2): 133-137, 14 figs.

. 1989. Ephydridae de la Republica Argentina

(Insecta Diptera). Serie Monografica y Didactica
No. 3. Facultad de Ciencias Naturales e Instituto Mi-
guel Lillo Universidad Nacional de Tucuman. 93 pp.
1990. Aportes al conocimiento del genero Beck-

eriella Williston, I. Descripcion de B. longivenths sp.
nov. y redescripcion de B. bispinosa (Diptera-Ephy-
dridae). Revista de la Sociedad de Entomologica Ar-
gentina, 48(1-4)1990(89): 143-147.

1991a. Aportes al conocimiento del genero

Beckeriella Williston. II. Descripcion de B. paragipas
sp. nov. y redescripcion de B. schildi. (Diptera-Ephy-
dridae). Acta Zoologica Lilloana, 40(1): 39-42.

1991b. Aportes al conocimiento del genero

Beckeriella Williston. III. Descripcion de B. pseudo-
clypeata, sp. n. y redescripcion de B. magnicornis
Cresson (Diptera, Ephydridae). Revista Brasileira de
Entomologia, 35(1): 45-48.

1992. Aportes al conocimiento del genero Beck-

Ephydridae). Revista Brasileira de Entomologia,
36(1): 229-233.

.1994. Aportes al conocimiento del genero Beck-

eriella Williston. Descripcion de B. filipina sp. nov.
(Diptera: Ephydridae). Neotropica, 40(103-104): 57-
59.

Mathis, W N. 1977. Key to the Neotropical genera of
Parydrinae with a revision of the genus Eleleides
Cresson (Diptera: Ephydridae). Proceedings of the Bi-
ological Society of Washington, 90(3): 553-565.

. 1986. Studies of Psilopinae (Diptera: Ephydri-
dae), I: A revision of the shore fly genus Placopsidella
Kertesz. Smithsonian Contributions to Zoology, 430:
1-30.

1990. A revision of the shore-fly genus Diphuia

eriella Williston. IV. Redescripcion de B. pendicornis
y descripcion de B. parapendicornis sp. n. (Diptera,

Cresson (Diptera: Ephydridae). Proceedings of the En-
tomological Society of Washington, 92(4): 746-756.

Mathis, W N, and T Zatwarnicki. 1995. A world cat-
alog of the shore flies (Diptera: Ephydridae). Memoirs
on Entomology, International, 4: vi+423 pp.

Mc Alpine, J. E 1981. Morphology and terminology
Adults, pp. 9-63. In McAlpine, J. F, et al., eds., Man-
ual of Nearctic Diptera. Ottawa. [Volume 1 is Mono-
graph 27 of Research Branch Agriculture Canada.]

Williston, S. W 1897. Diptera Brasiliana. Part IV Kan-
sas University Quarterly, series A, 6: 1-12.

Wirth, W W 1968. 77. Family Ephydridae, pp. 1-43.
In Papavero, N., ed., A catalogue of the Diptera of
the Americas south of the United States. Departamen-
to de Zoologia, Secretaria de Agricultura, Sao Paulo.

Zatwarnicki, T 1992. A new classification of Ephydri-
dae based on phylogenetic reconstruction (Diptera:
Cyclorrhapha). Genus, 3(2): 65-1 19.

MATHIS & GRIMALDI: SHORE FLIES BECKERIELLA

113

Chapter 5

The Tracheline Spider Genus Paccius
(Araneae, Corinnidae) in the Pare National de
Marojejy, Madagascar

Norman I. Platnick 1

Abstract

Five new species of the tracheline spider genus Paccius Simon are described from the Pare
National de Marojejy, Madagascar, including the first known females of the genus. Males have
extraordinarily modified setae on the palpal tibia that may serve to disperse pheromones or a
secretion that forms an epigynal plug in mated females. All five species found at Marojejy
appear to be endemic to the area as well as altitudinally segregated: P. angulatus at 450 m, P.
griswoldi and P. scharffi (belonging to different species groups) at 700-800 m, P. quinteri at
1625 m, and P. elevatus at 1875 m.

Resume

Cinq nouvelles especes d'araignee tracheline genre Paccius Simon ont ete decrites a partir
des specimens collectes dans le Pare National de Marojejy, Madagascar. Parmi elles se trouvait
le premier individu de sexe femelle connu appartenant a ce genre. Chez les males, les barbillons
du tibia sont extraordinairement modifies et sont remplaces par des poils raides qui doivent
leur servir pour disperser les pheromones ou la secretion qui forme le bouchon epigynal des
femelles avec lesquelles ils viennent de s'accoupler. Toutes les cinq especes trouvees a Marojejy
paraissent etre endemiques de l'aire en question de meme qu'elles semblent etre reparties selon
V altitude: P. angulatus a 450 m, P. griswoldi et P. scharffi (appartenant a differents groupes
d'especes) a 700-800 m, P. quinteri a 1625 m, et P. elevatus a 1875 m.

Introduction and Background

The spider genus Paccius Simon 1898 is an ob-
scure taxon. The type species was originally de-
scribed as Trachelas madagascariensis by Simon
(1889) on the basis of a single male, in poor con-
dition, from an unspecified locality on the island.
In his famous Histoire naturelle des araignees,
Simon originally (1897: 180) retained the species
in Trachelas L. Koch 1872, a genus that served
(then and now) as a "wastebasket" group for rel-
atively unmodified trachelines. In a supplement to

1 Division of Invertebrate Zoology, American Muse-
um of Natural History, Central Park West at 79th Street,
New York, NY 10024, U.S.A.

that consideration, however, Simon (1898a, p.
216) indicated that he had recently had the op-
portunity to study additional specimens, and es-
tablished the genus Paccius, with T. madagasca-
riensis as its type. Simon also included two ad-
ditional species, P. quadridentatus from the Sey-
chelles (subsequently described by Simon, 1898b,
p. 384) and P. mucronatus. Bonnet (1958, p.
3273) considered P. mucronatus a nomen nudum,
but Simon's (1898a) discussion provides putative-
ly diagnostic information (and even a type local-
ity, "Sainte-Marie de Madagascar"), and the
name is unquestionably available.

Simon published no illustrations of any of the
three species, however, and the genus has (per-
haps therefore) remained in obscurity. Benoit

PLATNICK: TRACHELINE SPIDER PACCIUS

115

(1978) reported on P. quadridentatus from the
Seychelles, illustrating a male and describing the
female, but his specimens are clearly just mis-
identified members of the widespread, synan-
thropic corinnid genus Oedignatha Thorell 1881.
In other words, males of three species of Paccius
were very briefly described a century ago, fe-
males of the genus remain unknown, and no il-
lustrations have appeared in the literature.
Through the courtesy of Dr. Christine Rollard of
the Museum National d'Histoire Naturelle
(MNHN), Paris, types of all three Simon species
have been available for comparison with the ma-
terial described below.

The absence of modern information on Paccius
is especially remarkable, for at least two reasons.
First, available collections, fragmentary as they
are, nevertheless indicate that the genus has ra-
diated extensively on Madagascar; there may be
as many as 25 species on the island (as well as
others from the Comoro Islands, La Reunion, and
Mauritius). All the species seem to be narrowly
endemic, so that the genus has great promise as a
subject for cladistic biogeography. Second, the
male palpal morphology is very unusual. The tib-
ial apophysis bears highly modified setae, one of
which is enormously enlarged and elaborated into
a scooped-out channel (Figs. 5-1, 5-2). Because
the channel appears to open into the interior of
the segment, it may be used for dispersing pher-
omones or the secretions that are used to produce
the epigynal plugs frequently found in (presum-
ably mated) females.

Here I describe just those specimens of Paccius
that have been taken in the Pare National (PN) de
Marojejy, Madagascar, where there appear to be
at least five species, all of which are new to sci-
ence and which are in large part altitudinally sep-
arated (the only two species that have been taken
at the same altitude, P. griswoldi and P. scharffi,
belong to different species groups within the ge-
nus). The collections studied include both those
of the recent expedition (currently housed in the
American Museum of Natural History, AMNH)
as well as those made by Dr. J. Coddington and
colleagues in 1993 and housed in the National
Museum of Natural History, Smithsonian Institu-
tion (USNM).

Paccius angulatus, new species

(Figures 5-1, 5-2)

Type Male holotype taken in a Malaise trap
at 450 m on a tributary of the Manantenina River,

8.0 km NW of Manantenina, in the Pare National
de Marojejy, 1426.2'S, 4946.5'E, Antsiranana
Province, Madagascar (13-24 October 1996; E.
Quinter, T Nguyen), deposited in AMNH.

Etymology The species name refers to the
shape of the base of the male retrolateral tibial
apophysis.

Diagnosis Males resemble those of P. ma-
dagascariensis but have a much smaller retrola-
teral expansion on the cymbium (Fig. 5-1), only
one (rather than two) modified tibial seta dorsal
of, and overlapping, the large channelized seta
(Fig. 5-2), and in having a dorsally expanded base
on the tibial apophysis itself (Fig. 5-2).

Male Total length, not including chelicerae,
8.6 mm. Carapace dark red, entire surface covered
with procurved rows of tubercles. From above,
both eye rows almost straight, posterior row wider
than anterior; from front, both rows slightly pro-
curved; anterior median eyes largest, posterior
medians smallest, lateral subequal; anterior me-
dians separated by less than their diameter, farther
from anterior laterals; posterior medians separated
by twice their diameter, by three times their di-
ameter from posterior laterals; lateral eyes of each
side separated by their diameter; median ocular
quadrangle wider than long, wider in back than
long. Clypeal height less than anterior median eye
diameter; chilum large, triangular, pointed and
protuberant medially. Chelicerae dark red, protu-
berant, with four teeth on each margin; retromar-
gin with lobe-shaped extension at base of fang.
Labium and endites dark red; labium narrowed at
about one-fourth its length, truncate distally; en-
dites depressed along medial edges but without
distinct median grooves. Sternum dark orange,
sides granulate, with triangular extensions to and
between coxae, not fused with epimeric sclerites,
which extend around fourth coxae, separating
them from pedicel.

Abdomen long, white, with orange epigastric
scutum encircling pedicel and occupying anterior
surface of dorsum; large, orange dorsal scutum
covering almost all of dorsum anteriorly, all of
dorsum posteriorly; venter with four irregular,
longitudinal rows of small, orange sclerites. Epi-
gastric scutum with arm-shaped posterolateral ex-
tensions extending around booklung openings,
which are bordered posteriorly by pair of rectan-
gular, sclerotic strips. Anterior lateral spinnerets
two-segmented, approximate, well removed from
tracheal spiracle; posterior median spinnerets
short, tubular; posterior lateral two-segmented.

Leg formula 1423; leg I orange, other legs yel-

116

FIELDIANA: ZOOLOGY

Figs. 5-1 to 5-4. 1, 2, Paccius angulatus, new species. 3, 4, P. elevatus, new species. 1, left male palp, ventral
view; 2, same, retrolateral view; 3, epigynum, ventral view; 4, same, dorsal view.

low; spines absent, metatarsi and tarsi I, II scop-
ulate, with two ventral rows of black denticles;
metatarsi III, IV with distal preening brushes; two
dentate claws accompanied by dense claw tufts;
trochanters unnotched.

Palpal tibia distally expanded, ventral portion
of expanded area unsclerotized, bearing three en-
larged setae; median seta boat-shaped, with inte-
rior channel; distal seta sinuous, crossing over
into channel of median seta at about half its
length; retrolateral tibial apophysis with large, ex-
panded, angular base (Fig. 5-2); cymbium with
retrolateral expansion occupying only about one-
fifth of cymbial length (Fig. 5-1); embolar base
slightly excavated.

Female Unknown.

Other Material Examined None.

Pare National de Marojejy, 1426.8'S, 4944.1'E,
Antsiranana Province, Madagascar (13-19 No-
vember 1996; E. Quinter), deposited in AMNH.

Etymology The specific name refers to the
relatively high altitude at which the species was
taken.

Diagnosis Females can easily be recognized
by the large lateral epigynal openings and elevat-
ed, longitudinal, median epigynal ridge (Fig. 5-3).

Male Unknown.

Female Total length, not including chelicerae,
8.8 mm. As in P. griswoldi, except for the follow-
ing. Cheliceral promargin with four or five teeth.
Epigynum with large, lateral openings and ele-
vated, longitudinal median ridge (Fig. 5-3); ducts
not curled posteriorly (Fig. 5-4).

Other Material Examined One female tak-
en with the holotype (AMNH).

Paccius elevatus, new species

(Figures 5-3, 5-4)

Type Female holotype taken beating vegeta-
tion at 1875 m at the source of the Andranomi-
fototra River, 1 1 .0 km NW of Manantenina, in the

Paccius griswoldi, new species

(Figures 5-5 to 5-8)

Types Male holotype and female allotype tak-
en on the forest floor at night at 700-800 m in

PLATNICK: TRACHELINE SPIDER PACCIUS

117

./Yj-i-M-r.'

1 *J\V'< \

Figs. 5-5 to 5-8. Paccius griswoldi, new species. 5, Left male palp, ventral view; 6, same, retrolateral view; 7,
epigynum, ventral view; 8, same, dorsal view.

the Pare National de Marojejy, 8.4 km NNW of
Manantenina, 1426'S, 4945'E, Antsiranana
Province, Madagascar (12 November 1993; C.
Griswold), deposited in USNM.

Etymology The specific name is a patronym
in honor of the collector of the types.

Diagnosis Males resemble those of P. ma-
dagascariensis but have a much smaller retrola-
teral expansion on the cymbium (Fig. 5-5) and
have the tip of the tibial apophysis well removed
from (rather than touching) the modified setae
(Fig. 5-6). Females resemble those of P. quinteri
in having long, straight median epigynal ducts,
but differ in having those ducts curled posteriorly
(Fig. 5-8).

Male Total length, not including chelicerae,
9.3 mm. As in P. angulatus, except for the fol-
lowing. Cheliceral promargin with five teeth. Ab-
dominal venter with longitudinal rows of sclerites
completely encased within large, orange ventral
scutum. All setae associated with retrolateral tibial
apophysis enlarged; median boat-shaped seta
crossed by two more distal setae, more proximal
setae forming thick white brush well separated

from digitiform tip of tibial apophysis (Fig. 5-6);
cymbium with retrolateral expansion occupying
only about one-fourth of cymbial length (Fig.
5-5); embolar base distinctly excavated.

Female As in male P. angulatus, except for
the following. Total length, not including chelic-
erae, 10.9 mm. Cheliceral promargin with five
teeth. Epigastric scutum restricted to sides and
venter; dorsal scutum restricted to small sclerite
above pedicel, on anterior surface of abdomen;
abdominal dorsum pale gray, with two pairs of
darker cardiac spots; venter pale gray, with two
longitudinal rows of few dark spots. Posterior me-
dian and posterior lateral spinnerets each with two
and one cylindrical gland spigots, respectively,
but posterior medians shaped as in male. Palpal
tibia and tarsus with elongated setae but without
spines; tarsal claw long, apparently smooth. Epi-
gynum with small hood and long, depressed atri-
um (Fig. 5-7); ducts recurved anteriorly, curled
posteriorly (Fig. 5-8).

Other Material Examined MADAGAS-
CAR: Antsiranana: PN de Marojejy, 8.4 km NNW
of Manantenina, 1426'S, 4945'E, 700-800 m,

118

FIELDIANA: ZOOLOGY

^K ^^ J/0:

Figs. 5-9 to 5-12. Paccius scharffi, new species. 9, Left male palp, ventral view; 10, same, retrolateral view; 11,
epigynum, ventral view; 12, same, dorsal view.

11-14 November 1993, beating foliage, on forest
floor and foliage at night (C. Griswold, N.
Scharff, USNM), 16", 4$; tributary, Manantenina
River, PN de Marojejy, 10.0 km NW of Manan-
tenina, 1426.0'S, 4945.7'E, 15-22 October
1996, beating vegetation, 750 m (E. Quinter, T.
Nguyen, AMNH), 1 9 .

Paccius scharffi, new species

(Figures 5-9 to 5-12)

Type Male holotype and female allotype tak-
en on foliage at 700-800 m in the Pare National
de Marojejy, 8.4 km NNW of Manantenina,
1426'S, 4945'E, Antsiranana Province, Mada-
gascar (12-14 November 1993; N. Scharff, J.
Coddington), deposited in USNM.

Etymology The specific name is a patronym
in honor of the collector of the holotype.

Diagnosis Males resemble those of P. mu-
cronatus in having a relatively large cymbial ex-
pansion (Fig. 5-9), but they can be distinguished
by the angular proximal corner of the cymbial ex-
pansion (Fig. 5-9) and the longer tip of the retro-

lateral tibial apophysis (Fig. 5-10). Females can
be recognized by the transverse epigynal hood
(Fig. 5-11) and anteriorly expanded epigynal
ducts (Fig. 5-12).

Male Total length, not including chelicerae,
6.8 mm. As in P. angulatus, except for the fol-
lowing. Abdominal venter with longitudinal rows
of sclerites completely encased within large, or-
ange ventral scutum. All setae associated with re-
trolateral tibial apophysis enlarged; median boat-
shaped seta crossed by two more distal setae,
more proximal setae forming thick white brush,
well separated from elongate, granulate tip of tib-
ial apophysis (Fig. 5-10); cymbium with retrola-
teral expansion occupying over half of cymbial
length, situated more distally than in other species
(Fig. 5-9); embolar base elongated, excavated.

Female Total length, not including chelicerae,
6.6 mm. As in P. griswoldi, except for the follow-
ing. Cheliceral promargin with four teeth. Epigyn-
um with small anterior pocket, transverse hood,
and wide, deeply depressed atrium (Fig. 5-11);
ducts thickened, recurved anteriorly, curled pos-
teriorly (Fig. 5-12).

Other Material Examined None.

PLATNICK: TRACHELINE SPIDER PACCIUS

119

M '4 // /

Figs. 5-13 to 5-16. Paccius quinteri, new species. 13, Left male palp, ventral view; 14, same, retrolateral view;
15, epigynum, ventral view; 16, same, dorsal view.

Paccius quinteri, new species

(Figures 5-13 to 5-16)

Types Male holotype and female allotype tak-
en in a yellow pan trap set at 1625 m along a
tributary at head of Andranomifototra River, 10.5
km NW of Manantenina, in the Pare National de
Marojejy, 1426.4'S, 4944.5'E, Antsiranana
Province, Madagascar (6-12 November 1996; E.
Quinter), deposited in AMNH.

Etymology The specific name is a patronym
in honor of the collector of the types.

Diagnosis Males resemble those of P. ma-
dagascariensis but have a much smaller retrola-
teral expansion on the cymbium (Fig. 5-13) and a
sharply narrowed tip of the retrolateral tibial
apophysis (Fig. 5-14). Females can be recognized
by the small anterolateral epigynal openings (Fig.
5-15) and long straight epigynal ducts (Fig. 5-16).

Male Total length, not including chelicerae,
8.0 mm. As in P. angulatus, except for the fol-
lowing. Abdominal venter with longitudinal rows
of sclerites completely encased within large or-
ange ventral scutum. All setae associated with re-
trolateral tibial apophysis enlarged; median boat-

shaped seta crossed by two more distal setae,
more proximal setae forming thick white brush,
well separated from subdistally sharply narrowed
tip of tibial apophysis (Fig. 5-14); cymbium with
retrolateral expansion occupying about one-fourth
of cymbial length (Fig. 5-13); embolar base ex-
cavated, recessed behind soft tissue of tibial tip.

Female Total length, not including chelicer-
ae, 5.3 mm. As in P. griswoldi, except for the
following. Cheliceral promargin with four teeth.
Only two or three denticles on metatarsi I, none
on tarsi I or leg II. Epigynum with small antero-
lateral openings (Fig. 5-15); ducts long, straight,
recurved anteriorly, not curled posteriorly (Fig.
5-16).

Other Material Examined None.

Acknowledgments

I thank J. Coddington for access to specimens,
M. Shadab for help with illustrations, and K. Cat-
ley and V. Ovtsharenko for helpful comments on
a draft on the text.

120

FIELDIANA: ZOOLOGY

Literature Cited

Benoit, P. L. G. 1978. Contributions a l'etude de la fau-
ne terrestre des lies granitiques de l'archipel des Se-
chelles (Mission P. L. G. Benoit J. J. Van Mol 1972).
Clubionidae, Ctenidae et Pisauridae (Araneae). Revue
de Zoologie Africaine, 92: 940-946.

Bonnet, P. 1958. Bibliographia Araneorum, tome II.
Douladoure, Toulouse, pp. 3027-4230.

Simon, E. 1889. Descriptions d'especes et de genres
nouveaux de Madagascar et de Mayott. Annales de la

Societe Entomologique de France, serie 6, 8: 223-
236.

. 1897. Histoire Naturelle des Araignees, tome 2.

Roret, Paris, pp. 1-192.
. 1898a. Histoire Naturelle des Araignees, tome

2. Roret, Paris, pp. 193-380.

-. 1898b. Arachnides recueillis en 1895 par M. le

Dr A. Brauer (de l'Universite de Marburg) aux Ties
Sechelles. Annales de la Societe Entomologique de
France, 66: 370-388.

PLATNICK: TRACHELINE SPIDER PACCIUS

121

Chapter 6

Fruit Flies as Ecological Indicators: Species Diversity
and Abundance of Drosophilidae (Diptera) Along an
Altitudinal Transect in the Pare National de
Marojejy, Madagascar

David A. Grimaldi, 1 Eric L. Quinter, 1 and Tain Nguyen 1

Abstract

Fifty-eight species of drosophilid flies were collected along an altitudinal transect ranging
from 450 m to 1875 m in the Pare National de Marojejy, northeastern Madagascar. Over a 6-
week period flies were collected at five stations (450, 750, 1225, 1625, and 1875 m) along the
altitudinal transect by means of Malaise traps, yellow pan traps, fruit-baited traps, and sweep-
ing. At least 70% of the species collected by each method were obtained exclusively by that
method, indicating that any one method will yield only a small fraction of the total fauna of
drosophilids. Greatest species diversity occurred between (and including) 750 m and 1625 m;
the proportion of species found at only one of the five stations was also highest between (and
including) 750 m and 1625 m. Thirty-six (62%) of the species were represented by five or
fewer specimens, which can seriously complicate analyses of relative abundance. However, the
faunal composition dramatically reflected altitude, with an average of 50% species replacement
between adjacent stations. Given the speciose nature of drosophilids (3,200 world species),
their ecological diversity, their abundance, and the abundance of cosmopolitan tramp species
in disturbed habitats, this is an extremely useful group to use in the rapid assessment of eco-
logical conditions and habitat differences.

Resume

Cinquante-huit especes de mouches drosophiles ont ete collectees le long d'un transect al-
titudinal allant de 450 m a 1875 m dans le Pare National de Marojejy, dans le Nord-Est de
Madagascar. Les mouches ont ete collectees dans cinq stations (450, 750, 1225, 1625, et 1875
m) pendant six semaines, le long d'un transect altitudinal, en utilisant des trappes "Malaise,"
des "yellow pan traps," des pieges appates avec des fruits et des balayures. 70% et plus des
especes collectees a partir de chaque technique n'etaient pas captures par les autres systemes
de piegeage, ce qui indique que n'importe laquelle de ces methodes contribuera a l'inventaire
d'une petite partie du total des drosophiles recenses. La plus haute diversite d'especes etait
trouvee entre 750 et 1625 m. Trente-six (62%) des especes sont representee par cinq specimens
ou moins ce qui complique serieusement les analyses au niveau de l'abondance relative. Ce-
pendant, la composition faunistique reflete dramatiquement 1' altitude, avec une moyenne de
50% de remplacement d'especes entre des stations adjacentes. Les drosophiles, avec une di-
versite specifique assez elevee (3,200 especes mondiales), une diversite ecologique et une

1 Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New
York, NY 10024-5192, U.S.A.

GRIMALDI ET AL.: FRUIT FLIES 123

abondance importantes ainsi qu'une abondance d'especes vagabondes cosmopolites dans les
habitats perturbes, constituent un groupe extremement utile pour un inventaire rapide des con-
ditions ecologiques et des differences pouvant exister entre les habitats.

Introduction

The family Drosophilidae is renowned largely
on the basis of a species that is one of the three
most intensively studied laboratory eukaryotes:
Drosophila melanogaster Meigen. As of 1986,
2,822 species had been described for the family
(Wheeler, 1981, 1986), and several hundred more
have been described since then. Species diversity
is overwhelmingly circumtropical, and hundreds
of species have yet to be formally described and
even discovered. Largely because of the frugivo-
rous habits of many species in Drosophila, for
which the family is best known, the actual eco-
logical diversity in the family is rarely appreci-
ated. Indeed, the drosophilids are among the most
ecologically diverse of all families of Diptera (re-
viewed by Ashburner, 1981). Species and entire
lineages exist that also breed: in fungi, including
dangerously toxic mushrooms (Jaenike et al.,
1983); as miners of stems and leaves; in living
flowers (Brncic, 1983); in the nephric exudates of
land crabs (Carson, 1974); as probable parasites
of spittle bug nymphs (Grimaldi & Nguyen,
1999); as inquilines in bee nests (Juillard, 1947,
1948); and as larval predators of black fly and
chironomid larvae and eggs (Tsacas & Disney,
1974), sessile homopterans (such as scale insects
and "white flies") (Grimaldi, 1988), spider eggs
(Swezey, 1929), and even the embryos of centro-
lenellid glass frogs (Grimaldi, 1994). Given such
species diversity and disparate ecology, Droso-
philidae would presumably have been used rou-
tinely for habitat assessment, but they have not,
perhaps because of the taxonomic skills required
for separating and identifying tropical species.

In October and November 1996, a team of bi-
ologists funded by the World Wide Fund for Na-
ture and the American Museum of Natural His-
tory (AMNH) explored the biodiversity of the
Pare National (PN) de Marojejy, with the aim of
determining biotic composition along an altitudi-
nal transect close to the highest summit in the
reserve. Besides various vertebrates and plants,
invertebrates studied included arthropods such as
spiders and various kinds of flies (see other chap-
ters in this volume). Faunistic treatment of Afro-
tropical drosophilids is quite advanced, based for-

tunately on the excellent descriptive taxonomy of
Tsacas, Chassagnard, and co-workers (i.e., Chas-
sagnard et al., 1997; Tsacas, 1990; Tsacas & La-
chaise, 1979; Tsacas et al., 1988). The drosophilid
fauna of Madagascar, however, is poorly known,
with only 37 species having been recorded as of
1981, of which 10 were known only from that
island (Tsacas et al., 1981). According to these
authors, 15 of the Malagasy species were shared
with mainland Africa, 8 with the Mascarene Is-
lands, and 5 with the Comoro Islands. Since the
publication of the Tsacas et al. 1981 review, sev-
eral expeditions by the Laboratoire de Biologie et
Genetique Evolutives of the Centre National de la
Recherche Scientifique (CNRS) and the Museum
National d'Histoire Naturelle (Paris) have been
conducted specifically to survey the Malagasy
drosophilids. Publication of the results of those
surveys is in progress (e.g., Chassagnard &
McEvey, 1992). Given that the much smaller and
more distant Mascarene Islands harbor 39 known
species, it would be reasonable to expect Mada-
gascar to have a fauna of 200 or more species.
Although the focus of our particular study is eco-
logical assessment, the results and specimens re-
sulting from this survey contribute to larger, long-
term systematic studies of the unique drosophilid
fauna of Madagascar.

Materials and Methods

Collection methods involved Malaise traps, yel-
low pan traps, fruit-baited Drosophila traps, and
sweep netting. For ideal or maximal comparisons
between collection sites, it is critical that the col-
lection techniques used be standardized as closely
as possible ("constant-effort trapping"; South-
wood, 1980). However, as thoroughly discussed
by Southwood (1980), this is rarely achievable.
Even the most seemingly constant-effort methods,
such as Malaise traps and yellow pan traps, differ
in their effectiveness, depending on variations in
the microhabitats in which they are placed. The
effectiveness of fruit baiting varies with micro-
habitat location and with the type, age, and quan-
tity of the bait. Nonetheless, all methods com-

124

FIELD1ANA: ZOOLOGY

bined give a reasonable approximation of propor-
tional differences in species abundance.

The Malaise traps were made from white "no-
see-um" netting, approximately 2 X 3 m in size,
and were customized for compact storage by use
of a "head" made of the netting, from which was
suspended a Whirl-pak bag of 80% ethanol (this
head obviates the use of larger, heavier plastic or
glass collecting heads). The netting head was
sprayed with a contact insecticide; when insects
that were funneled into the head touched the net-
ting, they dropped into the bag of alcohol below.
Malaise traps are a passive collection device that
intercept flying insects, particularly Diptera and
Hymenoptera (Southwood, 1980). Their effective-
ness is greatest at the edge of a forest, particularly
on a steep hill opposite or close to the canopy.
One or two Malaise traps were used at each sta-
tion and were left in place for the duration of the
survey. Each bag was emptied of its contents just
before we ascended to the next camp and on the
return trip to the base of the mountain.

Yellow pan traps are routinely used for collect-
ing parasitoid Hymenoptera (Southwood, 1980)
and are often used for some families of Diptera,
such as Empididae and Dolichopodidae. They
have never been used for collecting Drosophili-
dae, but they proved to be very useful here. Bright
yellow plastic bowls filled with soapy water were
placed in clusters of three at numerous favorable
sites (near streams, in mossy vegetated spots)
within each station or camp. These were emptied
every other day and the contents transferred to
alcohol. Yellow pan traps are an active collection
device, in that the yellow color appears to attract
certain insects, which then become caught in the
fluid.

Fruit-baited traps were based on a design first
developed by Grimaldi (1985) and used to study
seasonal changes in population densities of Dro-
sophila. The traps are made of two plastic, bowl-
like containers screwed together at the mouths.
The bottom container has holes in it for the flies
to enter; the interior of the bottom container is
painted black, to facilitate the flies' movement to
the upper container (drosophilids are phototactic).
The upper container has a fine-screened funnel in
it, and the mashed and yeasted bait is applied to
the sides of the container. Flies attracted to the
bait and the light cannot escape through the small
hole at the end of the mesh funnel. This device
actively collects large numbers of fruit-breeding
species, which are attracted to the aromatics pro-
duced by fermentation. Yeasted, mashed oranges

and/or bananas were used for bait, and the traps
were hung in shaded locations and emptied peri-
odically.

Sweep nets were successful only at the first two
stations because the terrain and understory vege-
tation at the higher sites prevented use of this
technique. It is a very effective method for col-
lecting drosophilid species not normally bait-
trapped, such as species that congregate on rotting
wood or in leaf litter.

Collection Sites

Collection sites were in areas surrounding each
camp. E. L. Quinter and T Nguyen did the field-
work at the 450 and 750 m stations; Quinter con-
tinued the fieldwork alone at the higher stations.
Floristic details of each station are provided in
Chapters 1 and 3 of this volume and are only sum-
marized here. Representative habitats at stations
2-5 are shown in Figs. 6-1 to 6-4.

Station 1 Elevation approximately 450 m,
location approximately 8 km NW of Manantenina
(GPS coordinates 1426.2'S, 4946.5'E). Surveys
conducted 5-13 October. The station was estab-
lished at the edge of a tributary of the Mananten-
ina River, surrounded on all sides by mountains
covered with dense tropical vegetation, some of
which consisted of adventitious species indicative
of former human occupation, mixed with the orig-
inal indigenous rain forest flora.

Station 2 Elevation approximately 750 m,
location approximately 10 km NW of Mananten-
ina (GPS coordinates 1426.0'S, 4945.7'E). Sur-
veys conducted 15-22 October. The station was
situated along a different tributary of the Man-
antenina River, again surrounded by rugged,
mountainous terrain, this time featuring sheer rock
faces as well as open areas in dense forest (Fig.
6-1) created by wet, moss-covered rock slides.

Station 3 Elevation approximately 1225 m,
location approximately 1 1 km NW of Mananten-
ina (GPS coordinates 1426.2'S, 4944.5'E). Sur-
veys conducted 25 October-3 November. The sta-
tion was located on a ridge surrounded by higher
ridges and peaks. Vegetation was considerably
different from that of the two previous sites, the
most conspicuous difference being the presence
of many scrubby trees luxuriantly encrusted with
lichens, especially trees along the ridgetops (Fig.
6-2).

Station 4 Elevation approximately 1625 m,
location approximately 10.5 km NW of Manan-

GRIMALDI ET AL.: FRUIT FLIES

125

Fig. 6-1. Habitat at 750 m station. (Photograph by Eleanor Sterling.)

tenina (GPS coordinates 1426.4'S, 4944.5'E).
Surveys conducted 6-12 November. The camp
was located in a swale between two ridges, along
a tributary at the head of the Andranomifototra
River. Vegetation at this site was similar to that at
the 1225 m station (Fig. 6-3).

Station 5 Elevation approximately 1875 m,
approximately 1 1 km NW of Manantenina (GPS
coordinates 1426.8'S. 4944.1'E). Surveys were
conducted 13-19 November. This station was
above timberline but approximately 250 m below
the summit of Marojejy. The camp itself was sit-
uated on a grass- and shrub-covered slope devoid

of all but a few shrubby trees encrusted with li-
chens (Fig. 6-4). The terrain was granitic, carpet-
ed by a very thick, spongy mat of moss, lichens,
and ericaceous shrubs, with very little soil. Even
the bamboos were dwarfed in size, being at most
2-3 m high. The summit was windy, rocky, and
barren.

Specimen Handling

Drosophilid specimens were stored in vials of
70% ethanol and were prepared at the AMNH.

126

FIELDIANA: ZOOLOGY

Fig. 6-2. Habitat (left) at 1225 m station, and Malaise trap in situ. (Photographs by Eleanor Sterling.)

They were critical point-dried and then mounted
onto pinned paper points (a standard procedure
for acalyptrate fly specimens); then they were
sorted carefully by genus and species on the basis
of external features of males and females. No dis-
sections were made; these will be done later, for
systematic reports in which more accurate species
identifications are required and new species will
be described. It is traditional in drosophilid sys-
tematics to base species definitions on males only;
as a result, female specimens are often ignored.
This approach does stabilize nomenclature, since
species are usually more easily separated on the
basis of males. However, as a result of this, a
plethora of female characters (e.g., Grimaldi,
1990; Grimaldi & Nguyen, 1999) have been ig-
nored. External features of colors and patterns,
wing vein ratios, number and position of aristal
branches, and other features often allow a reliable
association of males and females. We used the
classification of drosophilid genera presented by
Grimaldi (1990), in particular the placement of
Scaptodrosophila and Hirtodrosophila as separate
genera, not subgenera within Drosophila.

Results

Table 6-1 presents the data on numbers of in-
dividual flies of each sex collected at each station
and by each technique. They are identified as sp.
1, sp. 2, and so on, within each genus. The three
Zaprionus species, however, represent identifi-
able, described species as based on revisions of
this genus by Tsacas and Chassagnard (1990),
Chassagnard and McEvey (1992), Chassagnard
and Tsacas (1993), and other works by these au-
thors. Tables 6-2 and 6-3 summarize the species
diversity among sites and among trapping meth-
ods, respectively. Table 6-4 is a summary of the
number of species shared among the five altitu-
dinal stations.

Fifty-eight species of Madagascar Drosophili-
dae were collected (Table 6-1), based on 723 iden-
tified specimens, which is approximately 15 more
species than had been recorded from the island
thus far. This is also 25 more species than were
collected in a geographical transect across Kenya
by means of bait traps (Escher et al., 1997). Nev-
ertheless, the sampling for our survey was still

GRIMALDI ET AL.: FRUIT FLIES

127

Fig. 6-3. Malaise trap situated near 1625 m station, and in situ Drosophila bait trap. (Photographs by Eleanor
Sterling.)

128

FIELDIANA: ZOOLOGY

Fig. 6-4. In situ Malaise trap near 1875 m station. (Photograph by Eleanor Sterling.)

incomplete, as illustrated by the lack of specimens
for 12 genera represented by some 60 described
species in the African Region that are either re-
corded or suspected to be in Madagascar: Amiota,
Apenthecia, Cacoxenus, Erima, Gitona, Hirtodro-
sophila, Paraleucophenga, Dettopsomyia, Jen-
nealopsis, Lissocephala, Luzonimyia, and Zygo-
thrica. Some of these are restricted to specific
hosts, such as fleshy fungi {Hirtodrosophila and
Zygothrica) and decaying tree trunks {Amiota, Er-
ima, and Gitona). Some of these genera may be
restricted to the African mainland, such as
Jennealopsis and Apenthecia, the latter closely as-
sociated with Aloe and similar plants (Chassag-

nard et al., 1997). Unless one systematically
swept over blooms of fungi and up and down the
trunks of trees, large components of the fauna
would be (and apparently have been) missed. Less
explainable, however, is the absence from our col-
lections of Lissocephala, the virtual absence of
Leucophenga, and poor species representation of
Zaprionus, which are speciose genera in the Af-
rotropics. There are approximately 10 species of
Leucophenga in Madagascar alone and 60 species
from the mainland; they are presumably associ-
ated with, but almost certainly not restricted to,
fungi. Lissocephala has 17 Afrotropical species,
although none are apparently recorded from Mad-

GRIMALDI ET AL.: FRUIT FLIES

129

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130

FIELDIANA: ZOOLOGY

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GRIMALDI ET AL.: FRUIT FLIES

131

Table 6-2. Species Diversity Along Altitudinal Transect.

No. of

species at altitude

Variable

450 m

750 m

1225 m

1625 m

1875 m

No. of species

No. of exclusive species

Coefficient of Similarity*
Q

17
4

25
8

24
1

25
10

11
3

0.31
0.47

0.40
0.32

0.32
0.31

0.28
0.43

"Q = Jaccard's coefficient; C s = modified S0rensen's coefficient.

agascar; this genus breeds extensively in figs (Fi-
cus) (Lachaise, 1977; Lachaise & Tsacas, 1983).

Zaprionus has been the subject of revisionary
studies and currently consists of 53 species, with
14 species alone known from Madagascar (Chas-
sagnard & McEvey, 1992; Chassagnard & Tsacas,
1993). They are large, conspicuously colored flies
often abundant on decaying fruits, but only three
species were collected in bait traps in our study:
Zaprionus kolodkinae Chassagnard and Tsacas,
1987 {Zaprionus sp. 1 in Table 6-1); Z masca-
riensis Tsacas and David, 1975 (Zaprionus sp. 2);
and Z spinipilus Chassagnard and McEvey, 1992
(Zaprionus sp. 3). Zaprionus kolodkinae is en-
demic to Madagascar (throughout the country); Z
mascariensis occurs on Madagascar, the Comores,
Mauritius, and Reunion islands; and Z spinipilus
is known from Madagascar and the African main-
land (Cameroon and Malawi).

In contrast to the several aspects of incomplete
sampling in this study are several genera for
which our sampling is apparently better than
would be expected on the basis of the 1981 re-
view by Tsacas et al. For example, two species of
Chymomyza were found, where none were known
to exist from Madagascar; five Dichaetophora
species were collected, with only one previously
reported; 15 Scaptodrosophila species were col-
lected, with only three recorded from Madagascar
previously; four to five Liodrosophila (generic
identity of the fifth awaits confirmation), where
only three were known; and four Microdrosophila

were found where only one was previously re-
corded.

An unexpected result was the success of yellow
pan traps, which collected 108 specimens belong-
ing to 19 species; three-fourths of these species
were collected only in this manner. Groups col-
lected predominantly by yellow pan traps were
Dichaetophora, Liodrosophila, a species in or
near the genus Calodrosophila, and several spe-
cies of Drosophila s.s. (spp. 1, 2) and Scaptodro-
sophila (spp. 7, 8, 9, 11). The capture by Malaise
traps is as expected on the basis of previous ex-
periences in the Neotropics, with a respectable
species diversity based on relatively small sam-
ples. Fifteen species were collected with Malaise
traps, based on just 64 specimens. Groups col-
lected predominantly with Malaise traps were
Chymomyza and Neotanygastrella (they have sim-
ilar habits), several species of Scaptodrosophila
(sp. 1-3), the only species of Scaptomyza found,
and the four species of Microdrosophila. Approx-
imately three-fourths of the species found in Mal-
aise traps were collected only in this manner. The
number of specimens collected by hand netting is
meager (only 28) and reflects the inexperience of
the collectors; experienced collectors of droso-
philids usually acquire their most diverse samples
this way, relying on knowledge of breeding sites.
Interestingly, seven species were collected by
hand netting, five of which were acquired only in
this manner. Also expected is the abundance of
specimens collected with bait traps: 523 identified

Table 6-3. Drosophilid Diversity by Collection Method.

No. collected

by means of

Variable

Fruit bait traps

Malaise traps

Yellow pan traps

Netting

Total no. of species
No. (%) exclusive
No. of specimens

23

19 (82)
523

15

11 (73)

64

19

15 (78)
108

7

5(71)
28

132

FIELDIANA: ZOOLOGY

Table 6-4. Drosophilid Species Shared by Altitude.

Altitude

450 m

No. of shared species by altitude

750 m

1225 m

1625 m

1875 m

450 m

750 m

1225 m

1625 m

1875 m

17

10

8

2

1

25

14

4

2

24

12

5

25

8

11

specimens in 23 species, the highest diversity of
the four methods used. Since fruit-breeding spe-
cies tend to be abundant and ubiquitous, it is in-
teresting that 82% of the species found at fruit
baits were collected only in this way. Clearly, re-
lying on just one collection technique would give
a very incomplete sample of the fauna.

Species number did not vary significantly
among stations, the greatest difference being 11
species at 1875 m, with 17-25 species at the low-
er stations. Species composition, however, varied
dramatically. As shown in Table 6-2, the number
of species unique to a particular station was high-
est at 750 and 1625 m (eight and ten, respectively;
the 1225 m station was apparently anomalous,
with only one species unique to it). Two coeffi-
cients of similarity were used to quantify similar-
ities in species compositions: Jaccard's coefficient
and a modified S0rensen's coefficient (South-
wood, 1980). Jaccard's coefficient is defined as

Q'x = Ma + b) - j,

where j is number of species common to two sta-
tions and a and b are the total number of species
at each station. Unfortunately, this coefficient
gives equal weight to all species regardless of
abundance and rarity, and rare species can greatly
skew indices of similarity, since they are often
intercepted by chance. This is a particularly im-
portant consideration here, since 36 (62%) of the
drosophilid species were represented by five or
fewer specimens. Although a coefficient in which
similarity is weighted by the relative abundance
of shared species is preferable, it too is biased by
the sampling techniques used (an "active" meth-
od such as bait trapping gives a yield that is dis-
proportionately high compared with, say, Malaise
trapping). The modified S0rensen coefficient takes
relative abundance into account:

C N = 2, N /(aN + bN),

where aN and bN are the total numbers of indi-
viduals sampled in habitats a and b, and /N is the

sum of lesser values for species common to both
stations. As shown in Table 6-2, differences
among stations for the two indices is minor, the
most substantial difference being that the Jaccard
coefficient ranks the similarity between stations 2
and 3 as relatively high and the modified S0rensen
coefficient ranks the similarity between these two
stations as relatively low. This is probably be-
cause each of these stations has 12 species that
are represented by five or fewer specimens.

Table 6-4 reveals that, as expected, adjacent
stations have the most species in common. As one
goes to the next higher station, approximately half
the number of species, or fewer, remain in com-
mon. None of the stations can be identified as
more faunally distinct than another; they all show
a progressive and accumulated transformation of
the drosophilid fauna along the altitudinal transect
and are clearly distinct from the lowlands (<450
m) fauna. Since moisture, cold tolerance, and
plant hosts are all known to influence drosophil-
ids, all of these factors probably affect species
diversity and abundance of these flies.

An interesting observation is the absence of
cosmopolitan "tramp" drosophilid species in the
samples. This result stands in contrast to the find-
ing of cosmopolitan tramp noctuid moth species,
present at all stations (E. L. Quinter, pers. observ.)
but most abundant at the highest stations, where
the level of endemicity of the native noctuids is
also highest. This may reflect a greater vagility of
noctuids, combined with hilltopping behavior.
Species recognized for their ability to colonize
and thrive in areas highly disturbed by humans
are Drosophila melanogaster, D. simulans (these
two are native to central Africa). D. hydei (native
to the New World), D. immigrans (native to Asia),
D. funebris (native to Europe), D. busckii (native
to Asia), and most recently D. malerkotliana (na-
tive to Asia). These and other anthrophilic species
are highly tolerant of cool temperatures, desicca-
tion, and high concentrations of ethanol (a product
of fermentation in breeding sites), accept a range

GRIMALDI ET AL.: FRUIT FLIES

133

of hosts in which to breed, and have excellent
dispersal ability and reproductive capacity (Par-
sons & Stanley, 1981). Although some of our
bait-trapped species were abundant enough to be
considered tramps (e.g., D. melanogaster grp. spp.
14 and 16), a lack of the common cosmopolitan
species in our bait traps indicates that this area of
the PN de Marojejy is a relatively pristine envi-
ronment as yet unaffected by human encroach-
ment.

Acknowledgments

We are grateful to Steve Goodman (Field Mu-
seum of Natural History), Eleanor Sterling, and
Cal Snyder (American Museum of Natural His-
tory), who arranged the logistics for this field-
work. The fieldwork was sponsored by the World
Wide Fund for Nature and the AMNH Center for
Biodiversity and Conservation; the Center also
funded both the technicians who prepared speci-
mens and other aspects of the laboratory research
and the curation of specimens.

Literature Cited

Ashburner, M. 1981. Entomophagous and other bizarre
Drosophilidae. pp. 395-429. In Ashburner. M., et al.,
eds.. The Genetics and Biology of Drosophila, vol.
3a. Academic Press. London.

Brncic. D. 1983. The ecology of flower-breeding Dro-
sophila. pp. 333-382. In Ashburner. M., et al.. eds..
The Genetics and Biology of Drosophila, vol. 3e. Ac-
ademic Press. London.

Carson. H. L. 1974. Three flies and three islands: Par-
allel evolution in Drosophila. Proceedings of the Na-
tional Academy of Sciences, USA. 71: 3517-3521.

Chassagnard, M.-T. and S. E McEvey. 1992. The Za-
prionus of Madagascar, with descriptions of five new-
species (Diptera: Drosophilidae). Annals de la Societe
Entomologique de France, 28: 317-335.

Chassagnard. M.-T. and L. Tsacas. 1993. Le sous-
genre Zaprionus s.str. definition de groupes d'especes
et revision du sous-groupe vittiger (Diptera: Droso-
philidae). Annals de la Societe Entomologique de
France, 29: 173-194.

Chassagnard, M.-T, L. Tsacas, and D. L. Lachaise.
1997. Drosophilidae (Diptera) of Malawi. Annals of
the Natal Museum, 38: 61-131.

Escher. S. A., K. Eriksson, and G. Bachli. 1997. Col-
lection of Drosophilidae (Diptera) along a transect in
Kenya. Bulletin de la Societe Entomologique Suisse.
70: 1-14.

Grimaldi. D. 1985. Niche separation and competitive

coexistence in mycophagous Drosophila (Diptera:
Drosophilidae). Proceedings of the Entomological So-
ciety of Washington. 87: 498-51 1.

1988. Relicts in the Drosophilidae. pp. 183-

213. In Liebherr. J. K.. ed.. Zoogeography of Carib-
bean Insects. Cornell University Press. Ithaca.

1990. A phylogenetic. revised classification of

genera in the Drosophilidae (Diptera). Bulletin of the
American Museum of Natural History. 197: 1-139.
1994. Description and immature stages of Hi r-

todrosophila batracida sp.n. (Diptera: Drosophilidae).
a predator of frog embryos. Entomologica Scandinav-
ica. 25: 129-136.

Grimaldi. D.. and T. Nguyen. 1999. Monograph of the
spittle-bug flies, genus Cladochaeta (Diptera: Droso-
philidae: Cladochaetini). Bulletin of the American
Museum of Natural History. 241: 1-326.

Jaenike. J.. D. A. Grimaldi, A. E. Sluder. and A. L.
Greenleaf. 1983. Alpha-amanitin tolerance in my-
cophagous Drosophila. Science. 221: 165-167.

Juillard. C. 1947. Cacoxenus indagator Loew (Dipt..
Drosophilidae). Contribution a la biologie d*une par-
asite d'Osmia rufa L. Mitteilungen der Schweizerisch-
en Entomologischen Gesellschaft, 20: 567-593.

. 1948. Le comportement des larves de Ca-
coxenus indagator dans les nids de I'Osmia rufa. Mit-
teilunaen der Schweizerischen Entomologischen Ge-
sellschaft. 21: 547-554.

Lachaise, D. 1977. Niche separation of African Lisso-
cephala within the Ficus drosophilid community.
Oecologia. 31: 201-214.

Lachaise. D.. and L. Tsacas. 1983. Breeding sites in
tropical African drosophilids. pp. 221-332. In Ash-
burner. M.. et al.. eds.. The Genetics and Biology of
Drosophila. vol. 3e. Academic Press. London.

Parsons. P. A., and S. M. Stanley. 1981. Domesticated
and widespread species, pp. 349-393. In Ashburner
M.. et al.. eds.. The Genetics and Biology of Dro-
sophila. vol. 3a. Academic Press. London.

Solthwood. T. R. E. 1980. Ecological Methods. With
Particular Reference to the Study of Insect Popula-
tions, second ed. Chapman and Hall, London.

Swezey, O. H. 1929. Notes on the egg parasites of in-
sects in Hawaii. Proceedings of the Hawaiian Ento-
mological Society. 7: 282-292.

Tsacas. L. 1990. Drosophilidae de l'Afrique Australe
(Diptera). Annals of the Natal Museum. 31: 103-161.

Tsacas. L.. and M.-T. Chassagnard. 1990. Les especes
du genre Zaprionus a femur anterieurs spinuleux (Dip-
tera: Drosophilidae). Annals de la Societe Entomolo-
gique de France. 26: 461-487.

Tsacas. L.. and R. H. L. Disney. 1974. Two new Af-
rican species of Drosophila (Diptera. Drosophilidae)
whose larvae feed on Simulium larvae (Dipt.. Simu-
liidae). Tropenmedicin Parasitologic 25: 360-377.

Tsacas. L.. and D. Lachaise. 1979. La radiation afri-
caine des Lissocephala infeodes aux Ficus (Diptera:
Drosophilidae). Annals de la Societe Entomologique
de France. 15: 589-603.

Tsacas. L.. M.-T. Chassagnard. and J. R. David. 1988.
Un nouveau groupe d'especes afrotropicales antho-
philes dans le sous-genre Scaptodrosophila du genre

134

FTELDIANA: ZOOLOGY

Drosophila (Diptera: Drosophilidae). Annals de la So- Wheeler, M. R. 1981. The Drosophilidae: A taxonomic

ciete Entomologique de France, 24: 181-202. overview, pp. 1-97. In Ashburner, M, et al., eds., The

Genetics and Biology of Drosophila, vol. 3a. Aca-

Tsacas, L., D. Lachaise, and J. R. David. 1981. Com- demic Press, London.

position and biogeography of the Afrotropical droso- 1986. Additions to the catalogue of the world's

philid fauna, pp. 197-259. In Ashburner, M., et al., Drosophilidae, pp. 395-409. In Ashburner, M., et al.,

eds., The Genetics and Biology of Drosophila, vol. eds., The Genetics and Biology of Drosophila, vol.

3a. Academic Press, London. 3e. Academic Press, London.

GRIMALDI ET AL.: FRUIT FLIES 135

Chapter 7

Parasitic and Commensal Arthropods of Some
Birds and Mammals of the Pare National de
Marojejy, Madagascar

Barry M. OConnor 1

Abstract

Examination of 80 specimens representing 15 species of mammals and 8 specimens repre-
senting 7 species of birds collected in the Pare National de Marojejy yielded numerous collec-
tions of parasitic and commensal arthropods. All host species and all host individuals harbored
arthropod associates. Preliminary identifications of 17 families of mites (Acari) as well as
parasitic insects belonging to the orders Siphonaptera (fleas) and Phthiraptera (lice) are reported.
A species of the flower-inhabiting mite family Ameroseiidae is reported phoretic in the nares
of the yellow-bellied sunbird-asity, Neodrepanis hypoxantha, for the first time. A species of
Pseudoscorpionida is reported phoretic on Eliurus grandidieri, the first record of mammal-
phoretic pseudoscorpions from Madagascar. The ectoparasite fauna of the Pare National de
Marojejy is generally similar in its overall makeup to those of the Pare National d' Andringitra
and the Reserve Speciale d'Anjanaharibe-Sud.

Resume

L'examen de 80 specimens representant 15 especes de mammiferes et de 8 specimens re-
presentant 7 especes d'oiseaux, collectes dans le Pare National de Marojejy, a permis de dis-
poser de plusieurs collections d'arthropodes parasites et commensaux. Toutes les especes et les
individus hotes recelent des associations d'arthropodes. Des identifications preliminaires de 17
families de mites (Acari) ainsi que d'insectes parasites appartenant aux ordres de Siphonaptera
(puces) et de Phthiraptera (poux) sont rapportees. Une espece de mites qui habite les fleurs
(famille des Ameroseiidae) est rapportee d'etre phoretique dans les narines de Neodrepanis
hypoxantha (Philepitte faux-souimanga de Salomonsen) pendant un premier temps. Puis, une
espece de Pseudoscorpionida est rapportee d'etre phoretique sur Eliurus grandidieri, le premier
cas d' association mammiferes-pseudoscorpions enregistre a Madagascar. La faune ectoparasite
du Pare National de Marojejy est generalement similaire dans son ensemble avec celles du
Pare National d' Andringitra et de la Reserve Speciale d'Anjanaharibe-Sud.

Introduction agascar's unique vertebrate fauna on the basis of

specimens collected during faunal surveys led by

This chapter continues the study of parasitic Steven M. Goodman. I have previously reviewed

and commensal arthropods associated with Mad- the literature summarizing this parasite fauna and

provided preliminary lists of families and genera

1 Museum of Zoology, University of Michigan, Ann of mites and orders of insects associated with
Arbor, MI 48109, U.S.A. birds and mammals of the Reserve Naturelle In-

OCONNOR: PARASITIC AND COMMENSAL ARTHROPODS 137

tegrale cTAndringritra (subsequently reclassified
as a Pare National [PN]) (OConnor, 1996) and the
Reserve Speciale (RS) d'Anjanaharibe-Sud
(OConnor. 1998). The present contribution reports
the results of a similar survey conducted in the
PN de Marojejy in 1996.

Methods and Materials

A total of 80 specimens representing 1 5 species
of mammals and 8 specimens representing 7 spe-
cies of birds collected in the PN de Marojejy were
examined for arthropod parasites. Specimens were
collected by S. M. Goodman and his associates
during the survey of this area in 1996. Specimens
to be examined for parasites were collected and
processed, and information was entered into a da-
tabase as described in the previous studies (OCon-
nor, 1996, 1998).

The following list is a preliminary report of the
results of this sampling. Because final identifica-
tions and descriptions of new taxa for most col-
lections are still pending, I report here the para-
site/commensal faunas for each host species gen-
erally at the level of family or, occasionally, genus
or species. For each host species, the number of
individuals examined is indicated after the species
name, and the number of host individuals harbor-
ing a particular arthropod taxon is listed after the
taxon name if more than one host was examined.
Host specimens are vouchered in the Field Mu-
seum of Natural History (FMNH). Chicago;
FMNH museum catalog numbers for each speci-
men examined follow the host name. The bird
classification used here follows that presented in
Chapter 10.

Results

Class Mammalia

Order Lipotyphla
Family Tenrecidae

Microgale cowani (N = 7)
159522, 159529-31. 159533-4, 159659
Acari: Parasitiformes
Ixodidae (5)
Laelapidae (6)
Acari: Acariformes
Trombiculidae (7)

Myobiidae:

Microgalobia (3)

Madamyobia (2)
Glycyphagidae (3)
Atopomelidae:

Listrophoroides (Alistrophoroides) (7)

Listrophoroides (Madlistrophoroides) (6)

Microgale dobsoni (N = 1)
159541
Acari: Parasitiformes

Ixodidae
Acari: Acariformes
Trombiculidae
Atopomelidae:

Listrophoroides (Alistrophoroides)
Listrophoroides (Madlistrophoroides)

Microgale gymnorhyncha (N = 2)
159544, 159547
Acari: Parasitiformes
Ixodidae (1)
Laelapidae (1)
Acari: Acariformes
Trombiculidae (1)
Atopomelidae:

Listrophoroides (Alistrophoroides) (2)
Listrophoroides (Madlistrophoroides) ( 1 )

Microgale longicaudata (N = 1)

159546

Acari: Acariformes

Trombiculidae

Glycyphagidae

Atopomelidae:

Listrophoroides (A listrophoroides)
Listrophoroides (Madlistrophoroides)

Microgale monticola (N = 9)
159529. 159549-53, 159556, 159569, 159676
Acari: Parasitiformes
Ixodidae (5)
Laelapidae (9)
Acari: Acariformes
Trombiculidae (8)
Myobiidae:

Microgalobia (4)
Glycyphagidae (1)
Atopomelidae:

Listrophoroides (Alistrophoroides) (9)
Listrophoroides (Madlistrophoroides) (6)

Microgale parvula (N = 3)
159560-1, 159680
Acari: Parasitiformes
Ixodidae (2)

138

FIELDIANA: ZOOLOGY

Laelapidae (2)
Acari: Acariformes

Trombiculidae (3)

Glycyphagidae (1)

Atopomelidae:

Listrophoroides (Alistrophoroides) (2)
Listrophoroides (Madlistrophoroides) (3)

Sarcoptidae:
Notoedres (1)

Microgale principula (N = 1 )
159547
Acari: Acariformes
Myobiidae:

Microgalobia
Demodicidae
Trombiculidae
Atopomelidae:

Listrophoroides (Alistrophoroides)
Listrophoroides (Madlistrophoroides)
Sarcoptidae:
Notoedres (1)

Microgale soricoides (N = 3)
159554-5, 159564
Acari: Parasitiformes
Ixodidae (1)
Laelapidae (3)
Acari: Acariformes
Myobiidae:

Microgalobia (2)
Trombiculidae (2)
Atopomelidae:

Listrophoroides (Alistrophoroides) (2)
Listrophoroides (Madlistrophoroides) (3)

Microgale talazaci (N = 3)
159537, 159571, 159576
Acari: Parasitiformes
Ixodidae (2)
Laelapidae (2)
Acari: Acariformes
Trombiculidae (3)
Myobiidae:

Microgalobia (2)
Atopomelidae:

Listrophoroides (Alistrophoroides) (2)
Listrophoroides (Madlistrophoroides) (3)

Oryzorictes hova (N =
159578
Acari: Parasitiformes
Ixodidae:
Ixodes

Haemaphy sails
Laelapidae

1)

Acari: Acariformes

Glycyphagidae

Atopomelidae:

Listrophoroides (Alistrophoroides)
Listrophoroides (Madlistrophoroides)

Order Rodentia

Family Muridae

Eliurus grandidieri (N = 22)
Acari: Parasitiformes
Ixodidae (1)
Laelapidae (17)
Acari: Acariformes
Trombiculidae (22)
Demodicidae:

Demodex (3)
Glycyphagidae (1)
Atopomelidae:

Listrophoroides (Eulistrophoroides) (7)
Listrophoroides (Pallistrophoroides) (22)
Pseudoscorpionida

Probably Chernetidae (3)
Insecta: Phthiraptera: Anoplura

Unidentified family (19)
Insecta: Siphonaptera
Unidentified family (2)

Eliurus majori (N = 10)
159624-8, 159630, 159632, 159634, 159637-8
Acari: Parasitiformes
Laelapidae (7)
Pachylaelapidae (1)
Acari: Acariformes
Trombiculidae (10)
Demodicidae:

Demodex (1)
Atopomelidae:

Listrophoroides (Eulistrophoroides) (3)
Listrophoroides (Pallistrophoroides) (9)
Insecta: Phthiraptera: Anoplura
Unidentified family (8)

Eliurus minor (N = 2)
159639-40
Acari: Parasitiformes

Laelapidae (2)
Acari: Acariformes
Trombiculidae (2)
Demodicidae:

Demodex (2)
Atopomelidae:

Listrophoroides (Pallistrophoroides) (2)
Insecta: Phthiraptera: Anoplura

OCONNOR: PARASITIC AND COMMENSAL ARTHROPODS

139

Unidentified family (2)

Eliurus webbi (N = 5)
Acari: Parasitiformes
Ixodidae (1)
Laelapidae (4)
Pachylaelapidae (1)
Acari: Acariformes
Trombiculidae (5)
Demodicidae:

Demodex (2)
Glycyphagidae (2)
Atopomelidae:

Listrophoroides (Eulistrophoroides) (1)
Listrophoroides (Pallistrophoroides) (5)
Insecta: Phthiraptera: Anoplura
Unidentified family (1)

Voalavo gymnocaudus (2)
Acari: Parasitiformes

Laelapidae (2)
Acari: Acariformes
Trombiculidae (2)
Demodicidae:

Demodex (2)
Atopomelidae:

Listrophoroides (Eulistrophoroides) (2)
Listrophoroides (Pallistrophoroides) (2)

Class Aves

Order Passeriformes

Family Eurylaimidae

Neodrepanis coruscans (N = 1)
Acari: Acariformes
Proctophyllodidae
Trouessartiidae

Neodrepanis hypoxantha (N = 2)
Acari: Parasitiformes

Rhinonyssidae (1)

Ameroseiidae (1)
Acari: Acariformes

Proctophyllodidae (2)

Trouessartiidae (2)

Family Ploceidae

Foudia omissa (N = 1)

Acari: Acariformes

Proctophyllodidae

Avenzoariidae

Trouessartiidae

Family Pycnonotidae

Berneria zosterops (N = 1)
Acari: Acariformes
Analgidae
Proctophyllodidae
Avenzoariidae
Trouessartiidae
Insecta: Phthiraptera: Mallophaga
Unidentified family

1)

Family Timaliidae

Oxylabes madagascariensis (N
Acari: Acariformes

Analgidae

Proctophyllodidae

Avenzoariidae

Trouessartiidae
Insecta: Phthiraptera: Mallophaga

Unidentified family

Family Turdidae

Copsychus albospecularis (N = 1)
Acari: Acariformes
Trombiculidae
Ereynetidae
Analgidae
Proctophyllodidae
Trouessartiidae
Xolalgidae

Monticola sharpei (N =
Acari: Acariformes
Analgidae
Proctophyllodidae
Trouessartiidae

Discussion

1)

Comparison of the parasite fauna (compound
parasite communities) of the PN de Marojejy with
those of the PN d'Andringritra and RS
d'Anjanaharibe-Sud (OConnor, 1996, 1998) re-
veals a strong similiarity at the ordinal, familial,
and generic levels, suggesting that these parasite
communities are similar throughout the forested
regions of the country. The collection of a spec-
imen of the family Ameroseiidae from Neodre-
panis hypoxantha marks the first record of this
family in association with this group of birds.
Other ameroseiid species have been reported from
nectarivorous birds in Australia (Halliday, 1997).
Like other bird-phoretic ameroseiid mites, this

140

FIELDIANA: ZOOLOGY

specimen was recovered from the intranasal cav-
ity. Most flower-inhabiting Ameroseiidae disperse
on nectarivorous insects such as bees and butter-
flies. Haitlinger (1987) described Afrocypholae-
laps ranomafanaensis phoretic on an unidentified
butterfly from Madagascar. Phoretic pseudoscor-
pions are reported here for the first time from the
rodent Eliurus grandidieri. Several individuals of
this species harbored these symbionts; numerous
individuals were attached by their pedipalpal che-
lae to hairs of the hind legs. The pseudoscorpions
shared this attachment site with large laelapid
mites of the genus Androlaelaps. Since no other
mammalian host carried these organisms, some
specificity may be indicated.

Acknowledgments

I thank Steven M. Goodman for his extraordi-
nary efforts in making this study possible.

Literature Cited

Haitlinger, R. 1997. Afrocypholaelaps ranomafanaen-
sis sp. n. (Acari, Mesostigmata, Ameroseiidae) from
butterfly from Madagascar. Polskie Pismo Entomolo-
giczne, 57: 531-533.

Halliday, R. B. 1997. Revision of the Australian Ame-
roseiidae (Acarina: Mesostigmata). Invertebrate Tax-
onomy, 10: 179-201.

OConnor, B. M. 1996. Parasitic and commensal arthro-
pods of some birds and mammals of the Reserve Na-
turelle Integrale d'Andringitra, Madagascar, pp. 136
141. In Goodman, S. M., ed., A floral and faunal in-
ventory of the eastern slopes of the Reserve Naturelle
Integrale d'Andringitra, Madagascar: With reference
to elevational variation. Fieldiana: Zoology, n.s., 85:
1-319.

. 1998. Parasitic and commensal arthropods of

some birds and mammals of the Reserve Speciale
d'Anjanaharibe-Sud, Madagascar, pp. 73-78. In
Goodman, S. M., ed., A floral and faunal inventory
of the Reserve Speciale d'Anjanaharibe-Sud, Mada-
gascar: With reference to elevational variation. Fiel-
diana: Zoology, n.s., 90: 1-246.

OCONNOR: PARASITIC AND COMMENSAL ARTHROPODS

141

Chapter 8

Notes on a Small Collection of Fishes from the Pare
National de Marojejy, Northeastern Madagascar, with
a Description of a New Species of the Endemic Genus
Bedotia (Atherinomorpha: Bedotiidae)

Melanie L. J. Stiassny 1 and Ian J. Harrison 1

Abstract

A small collection of fishes from the Manantenina River in northeastern Madagascar is
reported. Included in the collection are six native species, two Malagasy endemics, and one
introduced species. One of the endemic taxa is a new species of the endemic rainbow fish
genus Bedotia. The new species, B. marojejy, described herein, is readily distinguished from
all congenerics by a characteristic pigmentation pattern and low vertebral count.

Resume

Une petite collection de poissons provenant de la Riviere Manantenina (nord-est de Mada-
gascar) est rapportee. Six especes natives, deux endemiques de Madagascar et une introduite
sont indues dans la collection. L'un des taxons endemiques est une espece nouvelle qui ap-
partient au genre Bedotia. La nouvelle espece, B. marojejy, decrite ici se distingue de ses
congeneres par des pigmentations caracteristiques et un faible nombre de vertebres.

Introduction

In a recent summary of the freshwater fishes of
Madagascar, Stiassny and Raminosoa (1994) pro-
vide an annotated listing of the native fishes of
the island. Although the continental waters of
Madagascar are relatively depauperate in terms of
overall species richness, there is a high-level of
endemism (84%) in the freshwater ichthyofauna.
Particularly striking is the fact that many of these
endemic taxa are basal representatives of their re-
spective clades (Stiassny, 1992; Stiassny & Ram-
inosoa, 1994). Despite the pronounced threat to
the island's freshwater ecosystems and the evo-

1 Department of Ichthyology, American Museum of
Natural History, Central Park West at 79th Street, New
York, NY 10024, U.S.A.

lutionary importance of many of these taxa, Mad-
agascar's freshwater fish fauna remains relatively
poorly documented and is often overlooked in
conservation efforts. Benstead et al. (2000) have
noted that if the criteria of the United States En-
dangered Species Act were applied to the fishes
of the island, 10 species would be considered crit-
ically endangered and 12 would be considered
threatened; insufficient data preclude accurate de-
termination of the status of most of the remainder.
There is clearly a strong impetus to continue with
ongoing attempts to assess the taxonomic com-
position, range, and conservation status of the
Malagasy ichthyofauna (Benstead et al., 2000;
Reinthal & Stiassny, 1991, 1997; Stiassny &
Raminosoa, 1994; Stiassny & Reinthal, 1992).

There have been few collections of fishes from
the region of the Pare National (PN) de Marojejy,

STIASSNY & HARRISON: FISHES

143

and with the exception of the description of a sin-
gle taxon, Rheocloides pelleghni Nichols and
LaMonte, 1931 (= Rheocles pellegrini; see Stiass-
ny, 1990), collected during the Mission Zoolo-
gique Franco- Anglo- Americaine (Rand, 1936)
from a locality "one day west of Andapa," no
records of freshwater fishes from this area occur
in the literature. In addition to the type of R. pel-
legrini, a number of other fishes collected during
the Mission Zoologique Franco-Anglo-Americai-
ne are deposited in the Ichthyology Department
of the American Museum of Natural History
(AMNH). and where appropriate these have been
consulted as comparative materials. So too has a
small collection of fishes from the Lokoho River,
downstream of Belaoko, made by Patrick De
Rham in October 1993 and deposited at the
AMNH.

During the 1996 faunal survey of the PN de
Marojejy a small collection of fishes from a lo-
cality at the banks of the Manantenina River at
the southeastern boundary of the reserve (Fig. 8-1),
about 28 km NE of the town of Andapa, was ob-
tained. In view of the general paucity of material
from this region and the pressing need for base-
line taxonomic and distributional data for Mala-
gasy fishes, a taxonomic report on the collection
is presented here. Included among the material is
a new species of rainbow fish of the endemic ge-
nus, Bedotia Regan, 1903, which is described
herein.

Abbreviations

AMNH, American Museum of Natural History;
BMNH, Natural History Museum, London;
MNHN, Museum National d'Histoire Naturelle.
Paris; RMNH. Nationaal Natuurhistorisch Muse-
um, Leiden; SL, standard length; TL, total length.

Taxonomic Review

1. Family Bedotiidae

The endemic Malagasy family Bedotiidae com-
prises the genera Bedotia Regan, 1903, and Rheo-
cles Jordan and Hubbs 1919. Although Rheocles
has been the subject of recent revision (Stiassny,
1990; Stiassny & Reinthal. 1992), the rainbow
fishes of the genus Bedotia have received consid-
erably less attention recently. Currently, the tax-

onomy of Bedotia is confused and greatly in need
of modern revisional study (Arnoult, 1959; Kie-
ner, 1963; Catala, 1975). We have begun this pro-
cess, and preliminary findings recognize five taxa
(B. madagascariensis Regan 1903, B. longianalis
Pellegrin 1914. and three undescribed species).
The specimens of Bedotia from the Manantenina
River are quite distinct from any of these other
nominal forms and are readily diagnosable on the
basis of a unique color pattern. Given the highly
distinctive nature of the Manantenina specimens
and the long-term nature of a comprehensive ge-
neric revision, the Manantenina species will be
described herein and subsequently incorporated
into a broader revisional study.

Bedotia marojejy, new species

(Figure 8-2)

Holotype AMNH 224647, female, 46.6 mm
SL. Banks of the Manantenina River (Lokoho
River basin) at the SE boundary of the PN de
Marojejy (approximate coordinates 1428'S,
4949'E), some 28 km NE of the town of Andapa.
Collected by local fisherwoman using traditional
dipnets. October-November 1996.

Paratypes AMNH 224648, two alcohol spec-
imens, one male, 47.7 mm SL, one undetermined
sex, 33.9 mm SL, one cleared and double-stained
specimen, undetermined sex, 45.3 mm SL. Same
data as holotype.

Diagnosis Bedotia marojejy is readily distin-
guished from all other nominal Bedotia by a dis-
tinctive color pattern characterized by a series of
dark blotches arrayed along the lateral midline. In
all other Bedotia species the lateral markings are
dominated by a broad midlateral band of pigmen-
tation, and similar blotching is never evident. B.
marojejy is further distinguished from other Be-
dotia by a low total vertebral count of 34 (includ-
ing the terminal, hypural-bearing half centrum),
compared to 35-37 (very rarely 34) in the other
species.

Description Morphological measurements
and meristic counts are given in Table 8-1. Mea-
surement of standard length, head length, snout
length, eye diameter, and caudal peduncle length
and depth follows Kottelat (1990). Body depth is
measured at the deepest part of the body, at the
origin of the pelvic fins. The distance from the
snout to the first or second dorsal fin is measured
straight from the tip of the snout to a vertical
through the base of the first fin ray in each fin

144

FIELDIANA: ZOOLOGY

\

s

\

\

Doany

Sambava

Farahalana

pahana

Fig. 8-1. Location of the PN de Marojejy (stippled area) in Madagascar. Arrow points to approximate locality in
which fishes were collected.

respectively. Anal length is measured from the
base of the first anal-fin ray to the base of the last
fin ray. Lower jaw length is measured from the
dentary symphysis to the posterior tip of the cor-
ner of the mouth. Currently the largest known
specimen of B. marojejy is a 47.7 mm SL male.

Bedotia marojejy are robust, relatively short-
bodied fishes somewhat deep-bodied anteriorly
and with moderate belly curvature. Dorsum of
head and nape broadly flatten to first dorsal fin.
Origin of first dorsal is well behind vertical
through pelvic-fin insertion, and origin of second
dorsal is well behind vertical through origin of
anal fin.

Snout is strongly indented behind prominent
premaxillary pedicels. Lower jaw is prognathous
and angled at about 30-35 to horizontal when
mouth is closed. Premaxilla and maxilla reach

vertical through anterior margin of orbit. Premax-
illae with deep lateral "bedotia notch" are char-
acteristic of the genus (Stiassny, 1990).

Teeth Anteriorly both upper and lower jaws
bear two to three rows of numerous small, strong-
ly recurved unicuspid teeth. The inner and outer
rows of teeth are poorly differentiated. The lower
jaw and the premaxilla posterior to the bedotia
notch each have a single row of teeth. A single
row of teeth is present along the anteroventral
face of vomer, and a tiny patch of endopterygoid
teeth is also present. No palatine or ectopterygoid
teeth are present, at least in individuals of sizes
available for examination.

Gill Rakers Two or three stout hypobranchial
rakers and eight or nine elongate ceratobranchial
rakers are present on the lower limb of the first
branchial arch. All rakers are strongly denticulate.

STIASSNY & HARRISON: FISHES

145

Fig. 8-2. Holotype of Bedotia marojejy sp. now. AMNH 224647. female. 46.6 mm SL.

Scales Body is covered with large, regularly
imbricate, cycloid scales. Predorsal scales number
16 or 17 along the dorsal midline. Thirty-two or
33 scales occur along the midlateral axis from just
behind the operculum, above the pectoral fin. to
the end of the hypural plate. Ten scales in trans-
verse series are located between the anal fin and
the second dorsal fin (including a very small scale
adjacent to each fin). One or two scales separate
the first and second dorsal fins. Twelve or 13
scales in transverse series are located entirely
around the caudal peduncle (circumpeduncular se-
ries). Dorsal, anal, and caudal scale sheaths and

axillary pelvic scales are absent. The body is fully
scaled.

Fins First dorsal fin bears six weak spines.
Second dorsal fin bears 12 or 13 rays, with the
first 3 or 4 unbranched. Anal fin bears 17 rays,
usually with the first 5 unbranched. Pectoral fin
bears 14 rays that are high set and relatively short,
the longest upper rays extending only a little be-
yond the vertical from the pelvic-fin insertion.
The pelvic fin has one weak spine and five strong-
ly bifurcate, branched rays. The caudal fin is
weakly emarginate.

Vertebrae Total vertebral count (taken from

Table 8-1. Bedotia

marojejy

. new

species.

Character Evaluated

Holotype

Paratypes

<N)

Min.*

Max.

Mean

Standard length

46.6

3

33.9

47.7

43.4

Head length "(^SL)

30.6

3

27.0

30.0

29.1

Snout length (% HL)

32.9

3

30.7

31.3

31.7

Eve diameter < 9t HL )

27.3

3

26.9

30.7

28.3

Body depth l r r SL)

yi >

3

23.7

24.0

23.7

Caudal peduncle length (9fc

HL)

54.5

3

54.6

57.5

55.5

Caudal peduncle depth (9

HL)

33.6

3

34.6

36.0

34.7

Snout first dorsal origin (

% SL)

54.4

3

52.1

52.8

52.9

Snout second dorsal origin ( r r SL)

65.3

3

63.9

65.4

65.0

Anal length (* SL)

28.3

3

27.0

29.7

28.1

Lower jaw length (9t HL)

51.0

3

48. ~

53.0

50.9

Character Evaluated

Holoty

pe

Paratypes

(N)

Range

Distribution

Longitudinal scales

33

3

7-)_

33

32 (1) 33 (3)

Gill rakers (lower arch)

11

3

11 (4)

Anal rays

17

3

17(4)

Second dorsal rays

12

3

12-

13

12 (3) 13(1)

Vertebrae total

34

3

34 (4)

(Precaudal - caudal -

terminal half centrum )

(17 - 16

- 1)

(17 - 16 - 1)(4)

''Min. and max. are the ranges only for the paratypical series, whereas the mean represents the entire sample.

146

FLFLDIANA: ZOOLOGY

radiographs and directly from the cleared and
double-stained specimen), 34: 17 precaudal ver-
tebrae (haemal arch not fused), 16 caudal verte-
brae (haemal arch fused), and terminal, hypural-
bearing half centrum. As noted in the diagnosis,
a total vertebral count of 34 is rarely encountered
in other Bedotia species. However, the sample
size for B. marojejy is small compared with the
number of specimens available for the five nom-
inal congeneric species, and intraspecific variation
within B. marojejy may be underestimated.

Preserved Coloration Ground color in both
male and female specimens is a pale creamy yel-
low. Dorsum of snout, nape, and body is darkly
pigmented. A black bar passes from the symphy-
sis of the lower jaw through the eye to the pos-
terodorsal border of the opercle. A dark crescentic
patch occurs at the pectoral-fin base. A series of
6 to 8 darkly pigmented blotches is arrayed along
the lateral midline. The ventrum of head, chest,
and body is pale creamy yellow. A broad wedge
of pigment occurring at the caudal-fin base ex-
tends as a narrow median stripe to the caudal mar-
gin. The caudal margin and ventral and dorsal
edges are darkly pigmented. The bases of the sec-
ond dorsal and anal fins are darkly pigmented, and
both fins have distal margins, also darkly pig-
mented. No data on live coloration are currently
available.

Viscera and Diet Gut extremely short, intes-
tinal length only about one-third body length. In
all four specimens the intestines are almost empty,
containing nothing but a few partially disarticu-
lated insects. In all individuals the remains of both
aquatic (larvae) and terrestrial insects were found,
suggesting that this species is probably essentially
insectivorous, opportunistically feeding on both
autochthonous and allochthonous sources.

Distribution Currently the species is known
only from the type locality in the Manantenina
River. In view of the paucity of collections from
this region, however, it is probable that additional
populations of B. marojejy are present in similar
habitats throughout the surrounding area.

Etymology The specific epithet was made in
reference to the locality in which the type series
was collected.

2. Family Anguillidae

Anguilla marmorata Quoy and Gaimard, 1824

(See Castle (1984) for synonymies)

One specimen of Anguilla marmorata was col-
lected at the Manantenina River site (AMNH

226649, 570 mm TL). It accords well with de-
scriptions of the species by Bauchot and Bianchi
(1984) and Skelton (1993) in having the dorsal-
fin origin nearer to the gill slit than to the anus,
the maxillary toothbands with distinctive longi-
tudinal grooves, and in being heavily mottled
brownish black. These features readily distinguish
A. marmorata from the three other anguillids cur-
rently recognized in Malagasy freshwaters: A.
mossambica (Peters, 1 852), A. bicolor McCleland,
1844, and A. bengalensis labiata (Peters, 1852).

Anguilla marmorata, commonly called the gi-
ant mottled eel, attains lengths of up to 185 cm
TL (Skelton, 1993). It is widespread in fresh and
brackish waters from sea level up to altitudes of
300-900 m throughout Madagascar (Kiener,
1963). Outside of Madagascar A. marmorata is
widespread from Western Cape Province to Ken-
ya in Africa (Skelton, 1993), the Comoros (Teu-
gels et al., 1985), and east to Polynesia and the
Ryukus, Japan.

Comparative Material Anguilla marmorata:
AMNH 88113, 460 mm TL, Toamasina (Tama-
tave), main marketplace in town center, Madagas-
car, 15-VII-1988, M. L. J. Stiassny, P. N. Reinthal;
AMNH 88125, 400 mm TL, Toamasina (Tama-
tave), main marketplace in town center, Madagas-
car, 15-VII-1988, M. L. J. Stiassny, P. N. Reinthal;
AMNH 58991, 62.4 mm, 170 mm, 175 mm TL,
creek crossing N coast road ca. 4.5 km N of Bo-
gia, Madang drainage, New Guinea, 19-X-1987,
L. R. Parenti, G. R. Allen, D. Coates.

Anguilla mossambica (Peters, 1852)

(See Castle (1984) for synonymies)

One specimen of Anguilla mossambica was
collected at the Manantenina River site (AMNH
226650, 320 mm TL). It too accords well with
descriptions of the species by Bauchot and Bian-
chi (1984) and Skelton (1993). Like A. marmo-
rata, A. mossambica has the dorsal-fin origin
nearer to the gill slit than to the anus, and this
readily distinguishes it from the other two Mala-
gasy anguillids. A. mossambica lacks the mottled
coloration, and its maxillary toothbands lack the
longitudinal grooves characteristic of A. marmor-
ata.

Anguilla mossambica, commonly called the Af-
rican longfin eel, attains lengths of up to 120 cm
TL (Skelton, 1993). It is the most widespread of
all Malagasy freshwater eels and is found from
sea level up to altitudes of over 900 m throughout

STIASSNY & HARRISON: FISHES

147

the island (Kiener, 1963). Outside of Madagascar
A. mossambica is commonly encountered on the
African mainland in east coast rivers from Kenya
south to Cape Agulhas, Republic of South Africa,
and it occurs on other western Indian Ocean is-
lands (Skelton. 1993).

Comparative Material Anguilla mossambi-
ca: AMNH 88114, 295 mm TL, Toamasina (Ta-
matave). main marketplace in town center. Mad-
agascar, 15-VII-1988. M. L. J. Stiassny. P. N.
Reinthal: AMNH 88010, 490 mm TL, town of
Ranomafana. Namorona River in shallow pools
cut off from mainstream by rocky outcrops, Mad-
agascar, 26-VI-1988, M. L. J. Stiassny, P. N. Rein-
thai; AMNH 11686, 399 mm TL, Madagascar.
1931. Archibold Expedition (= Mission Zoolo-
gique Franco-Anglo-Americaine). A. L. Rand. W.
H. du Mont.

3. Family Mugilidae

Agonostomus telfairii Bennett, 1831

(See Thomson (1986) for synonymies)

A single specimen of Agonostomus telfairii was
collected (AMNH 226651. 191 mm SL). Thom-
son (1997) recognized two Indian Ocean species
of Agonostomus in his revision of the Mugilidae.
namely A. catalai Pellegrin. 1932, with a thick
upper lip. and A. telfairii, with a thinner lip. In
the holotype of A. catalai, collected from the
Haute Mananano. Sahasinaka District. Madagas-
car, the mid-height of the upper lip is 2\% of the
head length: in the holotype of A. telfairii, col-
lected from Mauritius, the upper lip thickness is
10% of the head length. The specimen from the
Manantenina River has an upper lip thickness of
\1% head length. Examinations of comparative
material indicate a complete range of lip thick-
ness; for example, a collection from the Nosivolo
River (Mangoro drainage). Madagascar, includes
11 specimens of Agonostomus (AMNH 97100)
with lip thickness graded from 11% to 1 8% of
head length. Ingham (1952) and Thomson (1997)
stated that the thick-lipped species, A. catalai, could
be distinguished by having only a single row of
teeth in the lower jaw; however, collections ex-
amined in this study include specimens of Ago-
nostomus with thick lips and up to three irregu-
larly arranged rows of teeth in the lower jaw. Oth-
er distinguishing features for A. catalai given by
Thomson (1997) include the relative position on
the head of the upper part of the preorbital and

posterior tip of the upper jaw, relative position of
the pelvic fin. and number of gill rakers. However,
it is not possible to satisfactorily distinguish be-
tween the two species using these features.

Agonostomus telfairii (here including A. cata-
lai) is known from rivers of Madagascar, Anjouan
(Comoros), and the Mascarene islands of Reunion
and Mauritius (Thomson, 1980). According to Ca-
tala (1975), in Madagascar A. catalai (= A. tel-
fairii) is restricted to the upper reaches of fast-
flowing, rocky streams where it feeds exclusively
on algae. Kiener (1963) suggested that, in Mad-
agascar, A. telfairii would be unable to ascend the
rapids and waterfalls of the fast-flowing hill
streams.

Besides A. telfairii (including A. catalai), In-
gham (1952) and Thomson (1997) recognized
only one other species of Agonostomus, A. mon-
ticola. which is insectivorous and inhabits the up-
per reaches of torrential, rocky rivers in Central
and South America (Cruz, 1987; Phillip. 1993).
However. A. monticola is thought to migrate dur-
ing the rainy season down to coastal waters,
where it spawns before immediately returning up-
stream. After a short period at sea, the prejuven-
iles return to the rivers and migrate upstream
(Cruz. 1987; Ditty & Shaw. 1996). Hill streams
and rivers comprising isolated pools separated by
small waterfalls may not present a barrier to up-
stream migration because A. monticola is an ac-
complished leaper (K. Lazara, pers. comm.). A.
telfairii might show a similar, catadromous ecol-
ogy to A. monticola. A. telfairii can reach ca. 36
cm SL: it is taken in subsistence fisheries but not
commercially (Thomson, 1980).

The global distribution of Agonostomus is re-
markably disjunct; A. monticola is restricted to
rivers of Central America, northern South Amer-
ica, and the Galapagos Islands (Harrison, 1995,
Thomson. 1978, 1997), and A. telfairii is restrict-
ed to rivers of the aforementioned islands of the
western Indian Ocean. There is not sufficient in-
formation to provide any well-supported expla-
nation for this distribution of Agonostomus. How-
ever, two possible scenarios would benefit from
further analysis. One explanation is that this is the
relict of a former Tethyan distribution, with ex-
tension into the eastern Pacific during the Oligo-
cene and Miocene. This distribution would have
been disrupted by two vicariance processes: Mio-
cene closure of the Mediterranean, with the as-
sociated Messinian salinity crisis of that region,
and development of the current Panamanian isth-
mus during the Pliocene. A former Tethyan con-

148

FTELDIANA: ZOOLOGY

nection between Indo-Pacific and Atlantic rela-
tives has been discussed for various taxa; for ex-
ample, cods (Howes, 1991), gobies (Harrison,
1993; McKay & Miller, 1997), and anguillid eels
(Tagliavini et al., 1996; Tsukamoto & Aoyama,
1998). Anguillid eels, like species of Agonosto-
mus, have a catadromous lifestyle (McDowall,
1997). Vicariance biogeography of the Caribbean
and eastern Pacific regions has been discussed by
various authors (e.g., Coates & Obando, 1996;
Iturralde-Vinent & MacPhee, 1999; Rosen, 1975;
White, 1986).

An alternative hypothesis is that the current dis-
tribution of Agonostomus is a relict of a prolonged
Gondwanan link between Indo-Madagascar and
South America, via Antarctica, that may have last-
ed until ca. 80 million years ago (Sampson et al.,
1998). This scenario requires Agonostomus to be
at least twice as old as the oldest known fossil
mugilid, dated to 30-40 MYBP (Patterson, 1993).
This may be possible, considering information for
other smegmamorphs (sensu Johnson & Patterson,
1993). White (1986) discusses Cretaceous ances-
tral antherinids, although the earliest fossil mate-
rial is from the early Eocene, and the minimum
age of cyprinodontiforms is estimated to be late
Triassic, although the oldest described fossil cy-
prinodontiform is from the Oligocene (Parenti,
1981; Patterson, 1993). Contemporary distribu-
tion of Agonostomus through South America
would, according to this scenario, be related to the
very complex processes of Andean orogeny, as-
sociated changes in South American riverine
drainages, and periodic marine transgressions,
through the late Cretaceous and Tertiary (for dis-
cussion see Harrington, 1962; Lundberg, 1997;
Lundberg et al., 1998; Schaefer, 1997; Webb,
1995; Weitzman & Weitzman, 1982).

Comparative Material Agonostomus telfai-
rii: BMNH 1861.8.14.9, holotype, 108.5 mm SL,
Mauritius, Gerrard; AMNH 215499, 2 specimens,
79.3 mm and 84.7 mm SL, Lokoho River, Andapa
region, downstream Belaoko, northeast Madagas-
car, P. De Rham, 21-X-1993. Agonostomus telfai-
rii var. catalai: MNHN 1932-162, holotype, 129.8
mm SL, Haute Mananano, Madagascar, Catala;
BMNH 1865.9.21.4-5, 2 specimens, 175 and 195
mm SL, Johanna (?Anjouan); presented by Col.
Playfair; BMNH (no register no.), 126.5 mm SL,
Madagascar, Dr. S. E. Gray; AMNH 97100, 11
specimens, 136.5-227 mm SL, Tamatave Prov-
ince, Nosivolo River, below Zule's Village, deep
pools, Madagascar, M. L. J. Stiassny, P. N. Rein-
thai, and G. J. P. Naylor, 23-LX-1990; AMNH

971 12, 132 mm SL, Toamasina (Tamatave) Prov-
ince, Nosivolo River, 10 km upstream from Mar-
olambo, Madagascar, M. L. J. Stiassny, P. N.
Reinthal, and G. J. P. Naylor, 23-IX-1990. Nestis
cyphnoides: MNHN 5553, syntype, 207.5 mm
SL, lie de France (lie Maurice), Desjardins;
MNHN A.963, 2 syntypes, 163 mm and 146.2
mm SL, lie de France (lie Maurice), Dussumier;
MNHN A.4318, 2 syntypes, 67.8 mm and 65.8
mm SL, lie Bourbon (La Reunion), Leschenault.
Nestis dobuloides: MNHN A.4650, holotype, 360
mm SL, lie de France (lie Maurice), Lamarre Pi-
quot.

4. Family Cichlidae

Paratilapia polleni Bleeker, 1868

(See Marechal and Gosse (1991) for
synonymies)

Three specimens of the endemic cichlid genus,
Paratilapia, were collected at the Manantenina
River site (AMNH 226652; 54.7 mm SL, 50.6 mm
SL, 33.0 mm SL). Although most contemporary
authors follow Pellegrin's (1904) synonymy of
Paracara typus Bleeker, 1878, and Paratilapia
bleekeri Sauvage, 1891, with P. polleni Bleeker,
1868, several field biologists have noted the exis-
tence of distinct local phenotypes (Loiselle, 1995).
In a preliminary taxonomic note on the genus, Lo-
iselle and Stiassny (1993) resurrected Sauvage's P.
bleekeri, noting, among other features, the large
size of the spots and spangling of this species when
compared with the small spots of P. polleni. How-
ever, Loiselle and Stiassny (1993) accepted the
synonym Paracara typus with Paratilapia polleni,
mainly because of a paucity of comparative mate-
rial and the extremely poor state of the single type
specimen of P. typus. In the absence of a thorough
revision there remains considerable confusion re-
garding the species-level taxonomy of Paratilapia.
The problem of a definitive identification of the
Manantenina material is compounded by the fact
that the specimens lack the diagnostic coloration
and patterning of adult fish. However, based on a
morphometric appraisal, the Manantenina speci-
mens appear to approximate the more gracile, lon-
ger-jawed P. polleni phenotype than the deeper-
bodied, shorter-jawed P. bleekeri. Interestingly, al-
though the Manantenina specimens exhibit a typi-
cal juvenile Paratilapia patterning of 8-9 narrow,
closely spaced vertical bars and a large ocellated
tilapia-spot in the soft dorsal fin (Stiassny & Ger-

STIASSNY & HARRISON: FISHES

149

stner, 1992), the largest individual is a male with
advanced testicular development. According to
Kiener (1963), P. polleni are sexually mature at
about 12 cm on the Central High Plateau, com-
pared to 8 cm in the nutrient-poor acid waters of
the coastal plain where growth rates are retarded.
The Manantenina populations apparently are attain-
ing sexual maturity at a notably smaller size.

Prior to the extensive deforestation of the island
and the introduction of predatory exotic species,
Paratilapia polleni was the most widespread of
all Malagasy endemic cichlids, being recorded
from most regions of the island (Kiener, 1963;
Kiener & Mauge, 1966). However, because the
species cannot survive temperatures lower than
12-15C, it has always been absent from areas
much above 1500 m (Kiener, 1963). P. polleni has
been eradicated from much of its original range
by habitat alteration and introduced exotic spe-
cies. Today, although both P. polleni and P. blee-
keri are still widely distributed in eastern coastal
regions, they are nowhere abundant. P. polleni is
recorded as attaining sizes of 30 cm, although in-
dividuals of that size are rarely encountered today.

Comparative Material Paratilapia polleni:
RMNH 4.483, syntype, 119.3 mm SL, "Ambas-
suana" Madagascar septentrionalis in fluviis
(Nosy-be), 1864; RMNH 6.690, syntypes, 104.4
mm SL, 60.1 mm SL, "Ambassuana" Madagas-
car septentrionalis in fluviis (Nosy-be), 1864;
RMNH 3.994, syntype, 117.5 mm SL, "Ambas-
suana" Madagascar septentrionalis in fluviis
(Nosy-be), 1864; AMNH 18842, Region
d'Ankarana, Waterlot; AMNH 11689, Madagas-
car, Mission Zoologique Franco-Anglo-Americai-
ne (= Archibold Expedition), 28-IX-1931;
AMNH 97002, Bay Lake behind first dune, ca.
100 m from sea, east of road by Salehy Village,
1 km S of turnoff from Marolambo-Mananjary
Road, Toamasina (Tamatave) Province, Madagas-
car, 16-IX-1990, M. L. J. Stiassny, P. N. Reinthal,
G. J. P. Naylor; AMNH 97030, Mouth of Man-
goro River, opposite camp by Salehy Village, Toa-
masina (Tamatave) Province, Madagascar, 17-IX-
1990, M. L. J. Stiassny, P. N. Reinthal, G. J. P.
Naylor. Paracara typus: RMNH holotype, Mad-
agascar orientalis in fluviis (riviere de Mananara,
a 1' entree de la baie d'Antongil), specimen dis-
articulated. Paratilapia bleekeri: MNHN A.4195,
syntypes, 90.3 mm SL, 88.0 mm SL, 84.4 mm
SL, marais et rizieres de l'lmerina, pres
d'Antananarivo.

Oreochromis mossambicus (Peters, 1852)
(See Trewavas and Teugels (1991) for
synonymies)

Four juvenile specimens of the introduced Af-
rican cichlid Oreochromis mossambicus were col-
lected at the Manantenina River site (AMNH
226653; 63.6 mm SL, 56.2 mm SL, 52.6 mm SL,
34.1 mm SL). They conform to the descriptions
of the species (e.g., Trewavas, 1983; Skelton,
1993) and exhibit numerous vertical bars and a
characteristic series of 2-5 midlateral blotches
and some blotches of a more dorsal series. Ap-
parently O. mossambicus was introduced into
Madagascar from Mozambique in 1956 by the
Forestry Service for use in "rizipisciculture" (the
culture of fishes in flooded rice fields; Kiener,
1963). The species is now widespread throughout
the island.

5. Family Eleotridae

Eleotris fusca (Schneider, 1801)

(See Mauge (1986b) for synonymies)

Four specimens of Eleotris were collected at
the Manantenina River site (AMNH 226654; 2
males, 121.7 mm and 94 mm SL, and 2 females,
1 19.3 mm and 89 mm SL). They generally accord
with Akihito's (1967) and Hoese's (1986) descrip-
tions of E. fusca, having 8 or more gill rakers on
the first epibranchial, 61-66 scales in a longitu-
dinal series, infraorbital transverse rows 2, 4, and
6 penetrating below longitudinal row d, and 9
transverse infraorbital rows anterior to row b (the
last two characters are ambiguous in the smallest
specimen). These features distinguish E. fusca
from the very similar E. melanosoma Bleeker.
However, examination of comparative material in-
dicates that infraorbital papillae patterns are var-
iable. For example, specimens with high scale
counts (indicative of E. fusca) have an E. mela-
nosoma type papillae patterning, or intermediate
papillae patterns. It will not be possible to resolve
the taxonomy of Malagasy species of Eleotris
more precisely until a detailed review of the 13
nominal Indo-West Pacific species (Miller, 1998)
has been conducted.

Eleotris fusca attains 26 cm (Hoese, 1986). It
is currently reported as a widespread, epibenthic
species, found in fresh, brackish, and littoral wa-
ters throughout much of Madagascar and from the
east African coast eastward to Polynesia (Catala,

150

FIELDIANA: ZOOLOGY

1982; Mauge, 1986b). However, this distribution
requires confirmation through a taxonomic review
of the species.

Comparative Material Eleotris fusca:
AMNH 215521, male, 121 mm SL, Nosy-be, Dja-
bala River, near Ampombilava village, Madagas-
car, P. Loiselle, 18-IX-1990. Eleotris melanoso-
ma: AMNH 96904, male, 53.9 mm SL, Toama-
sina (Tamatave Province), main market place in
town center, caught nearby, Madagascar, local
fishermen, M. L. J. Stiassny, P. N. Reinthal, 15-
XII- 1988. Eleotris cf. melanosoma: AMNH
051509, male, 102.8 mm SL, 1 day W of Andapa,
northern Madagascar, Mission Zoologique Fran-
co- Anglo-Americaine (= Archibold Expedition),
A. L. Rand, W. H. du Mont, 2-X-1931; AMNH
097039, 7 specimens, 17.9-41.3 mm SL; Toa-
masina (Tamatave) Province, small affluent trib-
utary, River Mangoro, ca. 1 km upstream of Man-
goro ferry crossing, Madagascar, M. L. J. Stiass-
ny, P. N. Reinthal and G. J. P. Naylor, 17-LX-1990;
AMNH 97048, 1 male, 154 mm SL, 3 females,
64.6-102.2 mm SL, Toamasina (Tamatave) Prov-
ince, small stream ca. 20 km from Mangoro ferry
crossing on road to Marolambo, through banana
plantation, Madagascar, M. L. J. Stiassny, P. N.
Reinthal, and G. J. P. Naylor, 18-IX-1990.

the smaller male is slightly darker than the spe,
imen in photograph 10.

Awaous aeneofuscus inhabits fresh and brack-
ish waters of South Africa, Mozambique, Mada-
gascar, and the Comoros (Teugels et al., 1985;
Mauge, 1986a). Kiener (1963) reports that the
species may be found at altitudes up to 950 m. It
may attain 35 cm SL (Catala, 1982), although
larger specimens tend to be found not as far up-
river (Kiener, 1963).

Comparative material Awaous aeneofuscus:
AMNH 215496, 2 males, 85.8 mm and 91.2 mm
SL, Lokoho River, Andapa region, downstream of
Belaoko, northeast Madagascar, P. De Rham, 2-X-
1993; AMNH 88093, 1 male (from collection of
14 specimens), 134 mm SL, Pangalanes north of
Mangoro River, Mahanoro town behind Hotel de
la Pangalane, Madagascar, M. L. J. Stiassny, P. N.
Reinthal, 9-VII-1988; AMNH 097055, 3 females,
76.4-148 mm SL, Toamasina (Tamatave) Prov-
ince, Savalany River (small stream) by Ambodi-
sovoka village, bridge over road to Marolambo,
Madagascar, M. L. J. Stiassny, P. N. Reinthal, and
G. J. P. Naylor, 18-IX-1990.

Subfamily Sicydiinae

6. Family Gobiidae

Awaous aeneofuscus (Peters, 1852)

(See Mauge (1986a) for synonymies)

Two specimens of Awaous were collected at the
Manantenina River site (AMNH 226655; 1 fe-
male, 178 mm SL, and 1 male, 121.8 mm SL).
Both specimens conform to descriptions of A.
aeneofuscus (for example, Catala, 1982; e.g., with
26-36 small predorsal scales extending anteriorly
to a vertical through the preopercular canal, scales
absent on the opercle and preopercle, 55-60
scales in longitudinal series, and the mouth not
extending beyond the anterior of the eye. There
are two dark bands running anteroventrally from
the eye toward the mouth, and the body has a
series of dark brown patches just dorsal to the
midline (see Teugels et al., 1985, fig. 7). The pig-
mentation features are more distinct in the smaller
male specimen than in the large female. For a
general comparison, see photographs in Schmidt
et al. (1996, p. 34); the female appears similar to
the large specimen shown in photograph 2, and

Catala (1982) listed five sicydiine species from
Madagascar or neighboring islands, incorrectly
placing them all within the genus Sicydium Va-
lenciennes. These were: S. acutipinnis (Guiche-
not), S. franouxi Pellegrin, S. fasciatum Day, S.
laticeps Valenciennes, and S. lagocephalum (Pal-
las). Mauge (1986a) reassigned the species acu-
tipinnis Guichenot to its original genus, Cotylopus
Guichenot, 1863, and noted that reports of this
species from Madagascar are doubtful. Watson's
(1995) review of the genus and species concurred,
and he listed acutipinnis as known only from Re-
union. Additionally, Catala's (1982) description of
S. acutipinnis is inaccurate, citing a very high
scale count: 75 in longitudinal series, compared
to 46-63 cited by Watson (1995). The remaining
Malagasy species are all assigned to the genus
Sicyopterus Gill. 1861 (Mauge, 1986a). 5. laticeps
has been synonymized with S. lagocephalus
(Smith, 1959; Mauge, 1986a).

Sicyopterus fasciatus and S. franouxi are very
similar, and Mauge (1986a) listed Sicydium fra-
nouxi (= Sicyopterus franouxi) as a possible syn-
onym of Sicyopterus fasciatus. Pellegrin (1935)
and Catala (1982) refer to the absence of scales
on the median part of the nape of the neck and

STIASSNY & HARRISON: FISHES

151

median part of the abdomen in S. franouxi as a
character differentiating it from S. fasciatus. Ex-
amination of comparative material indicates that
differences in scalation of the nape are difficult to
detect and perhaps subjective. However, compar-
ison of similar-sized specimens of S. franouxi and
S. fasciatus (114 mm SL, AMNH 097149; and
115 mm SL, AMNH 215495, respectively) indi-
cates that the abdomen of S. fasciatus is entirely
scaled, whereas that of S. franouxi is less well-
scaled ventromedially. Smaller specimens of S.
franouxi also usually have the ventromedial part
of the abdomen less well-scaled, or naked, com-
pared to the ventrolateral parts of the abdomen,
although this is slightly variable. Pellegrin (1935)
referred to the coloration of S. franouxi as being
diagnostic, and this seems to be the most reliable
feature for differentiating it from S. fasciatus. In
most specimens of S. franouxi the anterior of the
snout, just above the upper lip, is dark brown and
continues as a dark brown band that runs beneath
the eye and over the mid part of the preopercle
towards its dorsoposterior margin. A prominent,
longitudinal dark brown band extends along the
midline of the body from the pectoral axilla to the
anterior half of the caudal fin. The rest of the body
is usually a lighter brown. This background pig-
mentation is darker in some specimens, however,
such that the longitudinal band, although still vis-
ible, is less distinct. Some of the lighter specimens
show traces of four or five dark brown, transverse
bars on the body, and Pellegrin's (1935) original
description refers to these. These transverse bands

may be more prominent in fresh specimens, as
suggested by a photograph, labeled as "Sicyop-
terus pranouxi" [sic] in Schmidt et al. (1996, p.
34, photograph 5). The base of the pectoral fin is
dark brown dorsally, at the same level as the lon-
gitudinal band on the body. The pectoral fin rays
are darkish brown near their bases but paler dis-
tally, producing a crescent-shaped band of dark
pigmentation over the fin. In comparative material
of Sicyopterus fasciatus the snout and sides of the
head are dark brown or dark gray, but this species
lacks the distinctive longitudinal band of dark
brown pigmentation that runs the length of the
body in S. franouxi. Day's (1876, p. 299) descrip-
tion and figure (pi. 54) of S. fasciatus give no
indication of a dark brown longitudinal bar. The
body coloration in S. fasciatus appears to be more
of a gray-brown than the light brown seen in S.
franouxi. The available comparative material of S.
fasciatus shows only vague traces of transverse
bands of dark brown pigment on the body, but
Day's description and figure indicate that the
bands are quite distinct. The pectoral fins of S.
fasciatus have a more uniform dark gray-brown
pigmentation than those of S. franouxi.

The possibility of conspecificity between S. fas-
ciatus and S. franouxi should not be discounted.
However, until a more detailed review of western
Indian Ocean sicydiines has been completed it
seems reasonable to distinguish between the S.
fasciatus and S. franouxi according to the squa-
mation and pigmentation features described
above. A provisional key for Malagasy species of
Sicyopterus follows:

la. Scales on nape and ventral part of abdomen much smaller than those on flanks; 58-63 scales in
longitudinal series (excluding scales on caudal fin) 2

lb. Scales on nape and ventral part of abdomen more or less equal in size to those on flanks; usually

52 or fewer scales in longitudinal series (excluding scales on caudal fin)

Sicyopterus lagocephalus

2a. Distinct, longitudinal, dark brown bar along midline of body; body otherwise lightish brown; pec-
toral fin darkish brown near base and on mid-part of fin but paler distally, producing a crescent-
shaped band of dark pigmentation over the fin; ventromedial part of abdomen usually lacking scales
Sicyopterus franouxi

2b. No dark brown longitudinal bar along midline of body; body grayish brown; pectoral fin uniform
dark gray-brown; ventromedial part of abdomen entirely scaled Sicyopterus fasciatus

Sicyopterus lagocephalus (Pallas, 1774)
(See Mauge (1986a) for synonymies)

One sicydiine collected at the Manantenina
River site (AMNH 226656, 1 female, 109.0 mm

SL) is identified as Sicyopterus lagocephalus ac-
cording to the above key. This species is darkish
gray-brown, with traces of five or six transverse,
darker brown, bands on the body and a longitu-
dinal, dark brown band running along the midline.

152

FIELDIANA: ZOOLOGY

The pectoral fins have a uniform dark brown pig-
mentation. The collected specimen (AMNH
226656) is slightly darker brown than other com-
parative material. Catala's (1982) scale count for
S. lagocephalus (57-65) is high according to the
available comparative material.

Sicyopterus lagocephalus inhabits swift-flow-
ing streams of Madagascar, Reunion, Mauritius,
and the Comoros (Mauge, 1986a; Teugels et al.,
1985; Watson, 1995). S. lagocephalus attains ca.
120 mm total length and is presumably amphidro-
mous like other sicydiine gobies; the adults re-
produce in freshwater, and the larvae are carried
down to the sea, where they remain for some time
before returning to freshwater to mature and re-
produce (Harrison, 1993; McDowall, 1997; Wat-
son, 1998). The ascent of enormous numbers of
postlarval S. lagocephalus up rivers of Reunion
and Madagascar has been reported by Valen-
ciennes in Cuvier and Valenciennes (1837) and
Kiener (1963). The migratory return of postlarvae
to freshwater more or less coincides with each
new moon; the more significant migrations in
Madagascar occur between April and September
(Kiener, 1963). The postlarvae are fished by local
people and eaten fresh or dried.

Comparative Material Sicyopterus lagoce-
phalus: AMNH 215498, 3 males, 76.4-104.0 mm
SL, Lokoho River, Andapa region, downstream of
Belaoko, northeast Madagascar, P. De Rham, 21-
X-1993. S.fasciatus: AMNH 215495, 1 male, 134
mm SL, and 1 female, 115.2 mm SL, Lokoho
River, Andapa region, downstream of Belaoko,
NE Madagascar, P. De Rham, 21-X-1993. S. fra-
nouxi: AMNH 097149, 4 females?, 77.1-113.6
mm SL, Toamasina (Tamatave) Province, Nosi-
volo River below Ambatomasina village, ca. 16
km E-NE of Marolambo, in large pool, Madagas-
car, M. L. J. Stiassny, P. N. Reinthal, and G. J. P.
Naylor, 21 -IX- 1990; AMNH 097071, 2 females,

52.6 mm and 83.6 mm SL, and one specimen of
undetermined sex, 54.6 mm SL, Toamasina (Ta-
matave) Province, Sahala River, near Andrano-
volo, Madagascar, M. L. J. Stiassny, P. N. Rein-
thal, and G. J. P. Naylor, 19-IX-1990; AMNH
097080, 4 females, 73.8-98.5 mm SL, 1 male,

63.7 mm SL, and 2 specimens of undetermined
sex, 67.8 mm and 95.5 mm SL, Toamasina (Ta-
matave) Province, Nosivolo River, below Ampa-
simaniona village, 26 km E-NE of Marolambo,
large side pool below rapids, Madagascar, M. L.
J. Stiassny, P. N. Reinthal and G. J. P. Naylor, 20-
IX-1990; AMNH 097068, 1 specimen of undeter-
mined sex, 38.6 mm SL, Toamasina (Tamatave)

Province, stream by Ambinanindrano, Madagas-
car, M. L. J. Stiassny, P. N. Reinthal, and G. J. P.
Naylor, 19-IX-1990.

Acknowledgments

Our thanks to Eleanor Sterling for her efforts
to obtain a sample of freshwater fishes during the
1996 expedition, and to Steve Goodman for his
invitation to prepare a short report on those fishes
for the present volume. Paul Loiselle (New York
Aquarium for Wildlife Conservation) kindly made
available some preliminary data on Bedotia tax-
onomy, and P. J. Miller (University of Bristol) and
F. Pezold (Northeast Louisiana University) pro-
vided some information concerning Eleotris, for
which we are grateful. Jean-Claude Razafimahai-
modison kindly helped locate some obscure lo-
calities. Our thanks also to Damaris Rodriguez for
all her help with the preparation of materials, data
collection, and x-rays, and to Erica Detwiler for
her excellent drawing of the holotype of Bedotia
marojejy. We are grateful to Herbert R. Axelrod
for his ongoing support for ichthyological re-
search.

Literature Cited

Akihito, Prince. 1967. On four species of the gobiid
fishes of the genus Eleotris found in Japan. Japanese
Journal of Ichthyology, 14(4/6): 135-166.

Arnoult, J. 1959. Faune de Madagascar. X. Poissons
des eaux douce. L'Institut de Recherche Scientifique,
Tananarive, 155 p.

Bauchot, M.-L., and G. Bianchi. 1984. Fiches FAO
d' identification des Guide des especes pour les be-
soins de la peches. Guides des poissons commerciaux
de Madagascar (especes marines et d'eaux saumatres).
Avec le support du Programme des Nations Unies
pour le Developpement (Projet RAF/79/065). Rome,
FAO, 135 p.

Benstead, J. P., M. L. J. Stiassny, P. V. Loiselle, K. R.
Riseng, and N. Raminosoa. 2000. River conservation
in Madagascar, pp. 203-229. In Boon, P. J., B. R. Da-
vies, and G. E. Petts, eds., Global Perspectives on Riv-
er Conservation: Science, Policy and Practice. Wiley,
New York.

Castle, P. H. J. 1984. Anguillidae, pp. 134-137. In Dag-
et, J., J. -P. Gosse, and D. F. E. Thys van den Aude-
naerde, eds., Check-list of the freshwater fishes of Af-
rica. Office de la Recherche Scientifique et Technique
Outre-Mer (ORSTOM), Paris and Musee Royal de
l'Afrique Centrale (MRAC), Tervuren.

Catala, R. 1975. Poissons d'eau douce de Madagascar.

STIASSNY & HARRISON: FISHES

153

Revue francaise d'Aquariologie et Herpetologie, 1:

29-32.

. 1982. Poissons d*eau douce de Madagascar. Re-

vue francaise d'Aquariologie et Herpetologie, 9(2):
57-64.

Coates, A. G., and J. A. Obando. 1996. The geologic
evolution of the Central American isthmus, pp. 21-
56. In Jackson. J. B. C, A. F. Budd. and A. G. Coates.
eds., Evolution and Environment in Tropical America.
University of Chicago Press, Chicago, IL.

Cruz, G. A. 1987. Reproductive biology and feeding
habits of cuyamel, Joturus pichardi and tepemechin,
Agonostomus monticola (Pisces; Mugilidae) from Rio
Platano, Mosquitia, Honduras. Bulletin of Marine Sci-
ence, 40(1): 63-72.

Cuvier, G., and A. Valenciennes. 1837. Histoire na-
turelle des poissons, 12. xxix + 507 pp., pi. 344-368.
Paris, Strasbourg.

Day, F. 1876. The fishes of India; being a natural history
of the fishes known to inhabit the seas and fresh wa-
ters of India. Burma and Ceylon. Part 2: 169-368. pi.
41-78. Wm. Dawson and Sons, London.

Ditty, J. G., and Shaw, R. F. 1996. Spatial and temporal
distribution of larval striped mullet (Mugil cephalus)
and white mullet (M. curema, family: Mugilidae) in
the northern Gulf of Mexico, with notes on mountain
mullet, Agonostomus monticola. Bulletin of Marine
Science, 59(2): 271-288.

Harrington, H. J. 1962. Paleographic development of
South America. Bulletin of the American Association
of Petroleum Geologists, 46(10): 1773-1814.

Harrison. I. J. 1993. The West African sicydiine fishes,
with notes on the genus Lentipes (Teleostei: Gobi-
idae). Ichthyological Exploration of Freshwaters, 4(3):
201-232.

. 1995. Mugilidae. pp. 1293-1298. In Fischer.

W., F Krupp, W. Schneider, C. Sommer, K. E. Car-
penter, and V. H. Niem, eds., Guia FAO para la iden-
tificacion de especies para los fines de la pesca. Pa-
cifico Centro-Oriental, 3. FAO, Rome.

Hoese. D. F 1986. Eleotridae, pp. 807-811. In Smith.
M. M., and P. C. Heemstra, eds.. Smith's Sea Fishes.
Springer- Verlag. Berlin.

Howes, G. J. 1991. Biogeography of gadoid fishes. Jour-
nal of Biogeography. 18: 595-622.

Ingham, S. E. 1952. The biology and taxonomy of the
Mugilidae. Unpublished manuscript in Library of Nat-
ural History Museum, London.

Iturralde-Vinent, M. A., and R. D. E. Macphee. 1999.
Paleogeography of the Caribbean region: Implications
for Cenozoic biogeography. Bulletin of the American
Museum of Natural History. 238: 1-95.

Johnson, G. D.. and C. Patterson. 1993. Percomorph
phylogeny: A survey of acanthomorphs and a new
proposal. Bulletin of Marine Science 52(1): 554-626.

Kiener, A. 1963. Poissons, peche et pisciculture a Mad-
agascar. Centre Technique Forestier Tropical, Nogent
sur Marne, 24: 244 p.

Kiener. A., and M. Mauge. 1966. Contributions a
l'etude systematique et ecologique des poissons Cich-
lidae endemiques de Madagascar. Memoires du Mu-

seum National d'Histoire Naturelle, Serie A, Zoolo-
gie, XL(2): 4-99.

Kottelat, M. 1990. Sailfin silversides (Pisces: Telma-
therinidae) of Lakes Towuti, Mahalona and Wawontoa
(Sulawesi, Indonesia) with descriptions of two new
genera and two new species. Ichthyological Explora-
tion of Freshwaters 1(3): 35-54.

Loiselle, P. V. 1995. The cichlids of Jurassic Park. Part
1. Cichlid News, 4(3): 18-23.

Loiselle, P. V., and M. L. J. Stiassny. 1993. How many
marakely? Journal of the American Cichlid Associa-
tion, 157: 2-8.

Lundberg, J. G. 1997. Freshwater fishes and their pa-
leobiotic implications, pp. 67-91. //; Kay, R. F. R. H.
Madden, R. L. Cifelli, and J. J. Flynn, eds.. Vertebrate
Paleontology in the Neotropics: The Miocene Fauna
of La Venta, Colombia. Smithsonian Institution Press,
Washington, D.C.

Lundberg. J. G.. L. G. Marshall, J. Guerrero, B.
Horton, M. C. Malabarba, and E Wesselingh.
1998. The stage for Neotropical fish diversification: A
history of tropical South American rivers, Chapter 1 .
In Malabarba. L. R.. R. E. Reis, R. P. Vari, C. A. S.
Lucena, and Z. M. S. Lucena, eds., Phylogeny and
Classification of Neotropical Fishes. Museu de Cien-
cias e Tecnologia, PUCRS. Porto Alegre, Brazil.

Marechal, C, and J. -P. Gosse. 1991. Paratilapia, pp.
358. In Daget. J., J. -P. Gosse. G. G. Teugels, and D.
F E. Thys van den Audenaerde, eds., Check-list of the
freshwater fishes of Africa. 4, Institut Royal des Sci-
ences Naturelles de Belgique (ISNB), Bruxelles, Mu-
see Royal de l'Afrique Centrale (MRAC), Tervuren,
Office de la Recherche Scientifique et Technique
Outre-Mer (ORSTOM). Paris.

Mauge, A. L. 1986a. Gobiidae, pp. 358-388. In Daget,
J.. J. -P. Gosse, and D. F E. Thys van den Audenaerde,
eds.. Check-list of the freshwater fishes of Africa. 2,
Institut Royal des Sciences Naturelles de Belgique
(ISNB). Bruxelles. Musee Royal de l'Afrique Centrale
(MRAC), Tervuren. Office de la Recherche Scienti-
fique et Technique Outre-Mer (ORSTOM), Paris.

. 1986b. Eleotridae. pp. 389-398. In Daget, J.,

J. -P. Gosse, and D. F E. Thys van den Audenaerde,
eds.. Check-list of the freshwater fishes of Africa. 2,
Institut Royal des Sciences Naturelles de Belgique
(ISNB), Bruxelles. Musee Royal de l'Afrique Centrale
(MRAC), Tervuren, Office de la Recherche Scienti-
fique et Technique Outre-Mer (ORSTOM), Paris.

McDowall, R. M. 1997. The evolution of diadromy in
fishes (revisited) and its place in phylogenetic analy-
sis. Reviews in Fish Biology and Fisheries, 7: 443-
462.

McKay. S. I., and P. J. Miller. 1997. The affinities of
European sand gobies (Teleostei: Gobiidae). Journal
of Natural History. 31: 1457-1482.

Miller. P. J. 1998. The West African species of Eleotris
and their systematic affinities (Teleostei: Gobioidei).
Journal of Natural History. 32: 273-296.

Parenti. L. R. 1981. A phylogenetic and biogeographic
analysis of cyprinodontiform fishes (Teleostei, Athe-
rinomorpha). Bulletin of the American Museum of
Natural History, 168(4): 335-557.

154

FIELDIANA: ZOOLOGY

Patterson, C. 1993. Osteichthyes: Teleostei, pp. 622-
656. In Benton, M. J., ed., The fossil record. 2. Chap-
man and Hall, London.

Pellegrin, J. 1904. Contribution a l'etude anatomique,
biologique et taxonomique des poissons des famille
des Cichlides. Memoires de la Societe zoologique de
France, 16: 41-402.

1935. Poissons de Madagascar recueillis par M.

Catala. Description d'un Sicydium nouveau. Bulletin
de la Societe zoologique de France, 60: 69-73.
Phillip, D. A. T. 1993. Reproduction and feeding of the
mountain mullet, Agonostomus monticola, in Trinidad,
West Indies. Environmental Biology of Fishes, 37:
47-55.

Rand, A. L. 1936. The distribution and habits of Mad-
agascar birds. Summary of the field notes of the Mis-
sion Zoologique Franco-Anglo-Americaine a Mada-
gascar. Bulletin of the American Museum of Natural
History, 72: 143-499.

Reinthal, P. N., and M. L. J. Stiassny. 1991. The fresh-
water fishes of Madagascar: A study of an endangered
fauna with recommendations for a conservation strat-
egy. Conservation Biology, 5(2): 231-243.

. 1997. Revision of the Madagascan genus Pty-

chochromoides (Teleostei: Cichlidae), with a descrip-
tion of a new species. Ichthyological Explorations of
Freshwaters, 7(3-4): 353-368.

Rosen, D. E. 1975. A vicariance model of Caribbean
biogeography. Systematic Zoology, 24(4): 431-464.

Sampson, S. D., L. M. Witmer, C. A. Forster, D. W.
Krause, P. M. O'Connor, P. Dodson, and F. Ra-
voavy. 1998. Predatory dinosaur remains from Mad-
agascar: Implications for the Cretaceous biogeography
of Gondwana, Science, 280: 1048-1051.

Schaefer, S. A. 1997. The Neotropical cascudinhos:
Systematics and biogeography of Otocinclus catfishes
(Siluriformes: Loricariidae). Proceedings of the Acad-
emy of Natural Sciences of Philadelphia, 148: 1-120.

Schmidt, W., and Aqua Geographia editors. 1996. La
Grande lie. Aqua Geographia, 13: 6-36.

Skelton, P. 1993. A Complete Guide to the Freshwater
Fishes of Southern Africa. Southern Books, South Af-
rica, 388 pp.

Smith, J. L. B. 1959. Gobioid fishes of the families Go-
biidae, Periophthalmidae, Trypauchenidae, Taenioidi-
dae, and Kraemeridae of the Western Indian Ocean.
Ichthyological Bulletin of Rhodes University, 13:
185-225.

Stiassny, M. L. J. 1990. Notes on the anatomy and re-
lationships of the bedotiid fishes of Madagascar, with
a taxonomic revision of the genus Rheocles (Atheri-
nomorpha: Bedotiidae). American Museum Novitates,
2979: 1-33.

-. 1992. Phylogenetic analysis and the role of sys-

tematics in the biodiversity crisis, pp. 109-120. In
Eldredge, N., ed., Systematics, Ecology and the Bio-
diversity Crisis. Columbia University Press, 220 pp.

Stiassny, M. L. J., and P. N. Reinthal. 1992. Descrip-
tion of a new species of Rheocles (Atherinomorpha:
Bedotiidae) from the Nosivolo tributary of the Man-
goro River, eastern Madagascar. American Museum
Novitates, 3031: 1-8.

Stiassny, M. L. J., and C. L. Gerstner. 1992. The pa-
rental care behaviour of Paratilapia polleni (Percifor-
mes, Labroidei), a phylogenetically primitive cichlid
from Madagascar, with a discussion of the evolution
of maternal care in the family Cichlidae. Environmen-
tal Biology of Fishes, 34: 219-233.

Stiassny, M. L. J., and N. Raminosoa. 1994. The fishes
of the inland waters of Madagascar. Biological diver-
sity in african fresh- and brackish water fishes. Geo-
graphical overviews. Annals Museum Royal Afrique
Centrale, Zoology, 275: 133-149.

Tagliavini, J., G. Gandolfi, A. M. Deiana, S. Salva-
dori. 1996. Phylogenetic relationships among two At-
lantic and three Indo-Pacific Anguilla species (Os-
teichthyes, Anguillidae). Italian Journal of Zoology,
63: 271-276.

Teugels, G. G., L. J. M. Janssens, J. Bogaert, and M.
Dumalin. 1985. Sur une collection de poissons de ri-
viere des Comores. Cybium, 9(1): 41-56.

Thomson, J. M. 1978. Mugilidae [unpaginated]. In Fi-
scher, W, ed., FAO species identification sheets for
fishery purposes. Western Central Atlantic (Fishing
Area 31), 3. FAO, Rome.

. 1980. Mugilidae [unpaginated]. In Fischer, W,

and G. Bianchi, eds., FAO species identification sheets
for fishery purposes. Western Indian Ocean (Fishing
Area 51), 3. FAO, Rome.

-. 1986. Mugilidae, pp. 344-349. In Daget, J., J.-

P. Gosse, and D. F. E. Thys van den Audenaerde, eds.,
Check-list of the freshwater fishes of Africa. 2, Institut
Royal des Sciences Naturelles de Belgique (ISNB),
Bruxelles, Musee Royal de l'Afrique Centrale
(MRAC), Tervuren, Office de la Recherche Scienti-
fique et Technique Outre-Mer (ORSTOM), Paris.

1997. The Mugilidae of the world. Memoirs of

the Queensland Museum, 41(3): 457-562.

Trewavas, E. 1983. Tilapiine fishes of the genera Sa-
rotherodon, Oreochromis and Danakilia. British Mu-
seum (Natural History), London, 583 p.

Trewavas, E., and G. G. Teugels. 1991. Oreochromis,
pp. 307-346. In Daget, J., J.-R Gosse, G. G. Teugels,
and D. F. E. Thys van den Audenaerde, eds., Check-
list of the freshwater fishes of Africa, 4, Institut Royal
des Sciences Naturelles de Belgique (ISNB), Bruxel-
les, Musee Royal de l'Afrique Centrale (MRAC), Ter-
vuren, Office de la Recherche Scientifique et Tech-
nique Outre-Mer (ORSTOM), Paris.

Tsukamoto, K., and J. Aoyama. 1998. Evolution of
freshwater eels of the genus Anguilla: A probable sce-
nario. Environmental Biology of Fishes, 52: 139-148.

Watson, R. E. 1995. Review of the freshwater goby
genus Cotylopus (Teleostei: Gobiidae: Sicydiinae).
Ichthyological Exploration of Freshwaters, 6(1): 61-
70.

. 1998. Stiphodon martenstyni, a new species of

freshwater goby from Sri Lanka (Teleostei: Gobiidae:
Sicydiini). Journal of Asian Natural History, 3(1): 69-
78.
Webb, S. D. 1995. Biological implications of the Middle
Miocene Amazon seaway. Science, 269: 361-362.

Weitzman, S. H., and M. Weitzman. 1982. Biogeo-
graphy and evolutionary diversification in Neotropical

STIASSNY & HARRISON: FISHES

155

freshwater fishes, with comments on the refuge theo- White. B. N. 1986. The isthmian link, antitropicality

ry. pp. 403-422. In Prance. G. T.. ed.. Biological Di- and American biogeography: Distributional history of

versification in the Tropics. Columbia University the Atherinopsinae (Pisces: Atherinidae). Systematic

Press. Zoology. 35(2): 176-194.

156 FIELDIANA: ZOOLOGY

Chapter 9

Herpetofaunal Species Diversity and Elevational
Distribution Within the Pare National de Marojejy,
Madgascar

A chille P. Rase lima nana, Christopher J. Raxworthy, 2 and
Ronald A. Nussbaum 3

Abstract

Amphibians and reptiles were surveyed along the summit trail of the Pare National (PN) de
Marojejy, between the village of Mandena, at 80 m elevation, and the Marojejy summit, at
2132 m elevation. A cumulative total of 75 days of surveying were completed during two
visits made in the rainy seasons of 1992 and 1996.

A total of 5 1 species of amphibians and 62 species of reptiles were found within the park,
and a further 3 amphibian and 7 reptile species were recorded within 5 km of the park boundary,
thus representing a regional herpetofaunal diversity of 123 species. Despite previous studies
made at Marojejy, 50% of the herpetofauna we recorded are new records for the park, and 10
species are undescribed. The actual herpetofaunal diversity of the PN de Marojejy is likely to
exceed 1 1 3 species as an additional 26 species are recorded from the park or region, thus
making this massif one of the most diverse areas of Madagascar. This exceptional species
diversity is almost certainly due to both the large elevational range found within the park and
the restricted elevational distribution of amphibians and reptiles in Madagascar. A weak mid-
elevation "bulge" in species diversity (possibly produced as an artifact of greater survey effort
at mid-elevation) was found at 700 m for both amphibians and reptiles, with minimum species
diversity occurring at the highest elevations in both groups. Endemism is also evident for the
Marojejy Massif, with 13 species currently known only from within the park.

The herpetofaunal species diversity and endemism of the PN de Marojejy demonstrate the
value of this park within the protected areas network on Madagascar. Future conservation
efforts must continue to maintain the integrity of the park, especially within its peripheral low-
elevation areas, which are most vulnerable to habitat loss.

Resume

Deux visites, d'une duree de 75 jours, le long du sommet du Pare National (PN) de Marojejy,
pendant les saisons de pluie des annees 1992 et 1996, ont permis de completer l'inventaire des
amphibiens et des reptiles entre le village de Mandena a 80 m jusque a la zone sommitale de
Marojejy a 2132 m.

Cet inventaire a permis de trouver 51 especes d' amphibiens et 62 especes de reptiles au sein
du pare, ainsi que 10 especes supplementaires aux alentours des limites du pare. Malgre les

1 WWF, B.P. 738, Antananarivo (101), Madagascar.

2 Division of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York,
New York 10024-5192, U.S.A.

3 Division of Herpetology, Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48103, U.S.A.

RASELIMANANA ET AL: HERPETOFAUNA 157

etudes effectuees a Marojejy auparavant, 50 pourcent de l'herpetofaune que nous avons enre-
gistres sont nouveaux pour le pare et 10 especes ne sont pas encore decrites. La diversite
actuelle de l'herpetofaune du PN de Marojejy est surement superieure a 113 especes, car un
supplement de 26 especes est enregistre pour le pare et les regions avoisinantes, faisant de ce
massif une des zones les plus riches en diversite de Madagascar. Cette diversite d'especes
exceptionnelle est surement due a la grande variation de 1' altitude au sein du pare et a la stricte
distribution altitudinale des amphibiens et reptiles de Madagascar. A 700 m d'altitude, il y a
eu un de diversite a moyenne altitudinale pour les amphibiens et les reptiles, avec un minimum
de diversite aux plus hautes altitudes pour les deux groupes. L'endemicite est aussi evidente
pour le Massif de Marojejy avec 13 taxa connus pour le pare seulement.

La diversite et l'endemicite de l'herpetofaune du PN de Marojejy montre la valeur de cette
reserve au sein du reseau des aires protegees de Madagascar. Les efforts de conservation
doivent continuer afin de maintenir l'integrite du pare, particulierement les zones peripheriques
de basse altitude qui sont les plus vulnerables a la perte d'habitat.

Introduction

The Pare National (PN) de Marojejy (previous-
ly classified as the Reserve Naturelle Integrate
[RNI] no. 12 de Marojejy or Marojezy in older
publications; see Chapter 1), with an area of
60,050 ha, includes the entire Marojejy Massif,
ranging between 75 and 2132 m elevation. The
massif is in northeastern Madagascar, with its
summit at 1424.03'S, 4944.00'E. The primary
vegetation of the park is rain forest, with ericoid
heathland at elevations above 1800 m (Nicoll &
Langrand, 1989; see Chapter 3). With the excep-
tion of the RNI de Tsaratanana, PN de Marojejy
includes the largest elevational range within any
protected area of Madagascar.

Although the Marojejy summit was first visited
in 1937 by the Service Geographique, the herpeto-
fauna of the massif remained unknown until collec-
tions were made in 1968 and 1972 by Charles P.
Blanc and a multidisciplinary French research team
(see Guillaumet et al., 1975). The 1968 collection
resulted in the descriptions of Brookesia karchei and
Calumma gastrotaenia tnarojezensis, the first rep-
tiles described from the massif (Brygoo et al.,
1970a,b). The second collection, 4 years later, led to
the description of another three chameleon taxa
(Brygoo et al., 1974a,b). Snakes collected during
these surveys have been described more recently by
Domergue (1984, 1988), and the Marojejy amphib-
ians collected by Blanc were listed by Blommers-
Schlosser and Blanc (1991).

In 1988, a British student expedition conducted a
survey of birds of the PN de Marojejy that also
included observations for 1 7 species of reptiles (Saf-
ford & Duckworth, 1990). However, because no

reptile vouchers were taken, tentative identifications
were based on color photographs. The first amphib-
ian and reptile inventory list for the park was pub-
lished by Jenkins (1987), who used published liter-
ature taken from Blanc's collections. This list was
further expanded by Nicoll and Langrand (1989) on
the basis of additional literature sources and proba-
bly some personal observations (without vouchers)
made by the authors during unspecified visits to the
park prior to 1989. Nicoll and Langrand (1989) list-
ed 17 amphibian and 22 reptile species.

The most recent herpetological inventory for
Marojejy was published as localities within spe-
cies accounts by Glaw and Vences (1994); it is
based on a survey of the literature and at least one
visit to the park made by one of the authors prior
to the publication of their book. The dates of the
survey are not given in this publication, but
Vences et al. (1997) describe two visits made by
Glaw to Marojejy, one in March 1994 and the
second in February 1995. A total of 50 amphibian
and 24 reptile species are listed from Marojejy by
Glaw and Vences (1994).

In this chapter we report the results of our own
surveys of the PN de Marojejy herpetofauna,
completed during two surveys of the park. The
first survey (University of Michigan) was made
between 16 November and 5 December 1992, and
the second survey (American Museum of Natural
History/Universite d' Antananarivo/World Wide
Fund for Nature) was made between 4 October
and 19 November 1996. The aim of these studies
was to survey the complete elevational range
within the park, to describe the distribution and
diversity of amphibians and reptiles within the
elevational gradient of the massif.

158

FIELDIANA: ZOOLOGY

Study Sites

The following five sites (all positioned along
the trail between the village of Mandena and the
Marojejy summit) were surveyed during 1992 and
1996.

Site 1 Ampanasankolana (Bedinta), by the Am-
binanitelo River, and Ampanasatongotra, 8 km
NW of Manantenina Village, by the Mananten-
ina River; 1426.2'S, 4946.5'E. Elevation 300-
525 m. Dates: 16 November and 3-5 December
1992, 4-13 October 1996.

Site 2 Andampimbazaha Cascade, on the Am-
bavaomby River, a tributary of the Manantenina
River; 1426.0'S, 4945.7'E. Elevation 550-850
m. Dates: 16 November-2 December 1992, 14-
24 October 1996.

Site 3 Antranohofa, by the Ambavaomby Ridge;
1426.2'S, 4944.5'E. Elevation 1050-1350 m.
Dates: 27-30 November 1992, 24 October-3
November 1996.

Site A Along the tributary Andranomifotrotra
River; 1426.4'S, 4944.5'E. Elevation 1550-
1700 m. Dates: 4-12 and 19 November 1996.

Site 5 Lohan Andranomifotrotra, Marojejy sum-
mit; 1426.4'S, 4944.1'E. Elevation 1800-
2133 m. Dates: 30 November 1992, 13-19 No-
vember 1996.

Methods

The members of the herpetological field survey
team during the 1992 survey were Ronald A.
Nussbaum, Gabriella Raharimanana, Christopher
J. Raxworthy, Angelin Razafimanantsoa, and An-
geluc Razafimanantsoa; those during the 1996
survey were Achille P. Raselimanana and Bruno
Zeze. The surveys were done during the early to
middle period of the rainy season (October-De-
cember), when herpetofaunal activity is at its
highest. Field techniques used to sample animals
(by both day and night) included (1) pitfall trap-
ping with drift fences; (2) visual and acoustic
searching; and (3) refuge examination (under and
in fallen logs and rotten tree stumps; under bark;
under rocks; in leaf litter, root- mat, and soil; and
in leaf axils of Pandanus screw palms and Rav-
enala traveller's palm).

The pitfall traps were buckets (275 mm deep,
290 mm top internal diameter, 220 mm bottom
internal diameter) with the handles removed and

small holes (2 mm diameter) punched in the bot-
tom to allow water drainage. Buckets were sunk
into the ground below a drift fence made from
plastic sheeting (0.5 m high) stapled in a vertical
position to thin wooden stakes, with the fence bot-
tom sealed 50 mm deep in the ground with soil
and leaf litter. The traplines were checked each
morning and late afternoon. After rain the buckets
were sponge-dried. The drift fence (100 m in
length) was positioned to run across the middle
of each pitfall trap. Pitfall traps were positioned
at both ends of the drift fence, with the other nine
traps at 10 m intervals. Three lines were used,
placed in the following forest types: ridge (along
the crest of a ridge), slope (on a gradient, inter-
mediate between ridgetop and valley bottom), and
valley (within 20 m of a stream in a valley bot-
tom). This is the procedure used in previous sur-
veys (e.g., Raxworthy & Nussbaum, 1994, 1996a)
and is recommended for use in future projects, to
allow standardization of capture techniques and
subsequent comparisons of results.

Visual searches and refuge examinations were
made throughout the full elevation range of habitats
available in the park. Night searches using head-
lights were made. The following information was
recorded for each individual at the time of capture:
date, time, longitude and latitude, elevation (mea-
sured by means of altimeters calibrated to 1 :
100,000 topographic maps), microhabitat, and cir-
cumstances of capture. Animals not retained as
specimens were released at the site of original cap-
ture. Voucher specimens were selected on the basis
of the following criteria: (1) documentation of spe-
cies diversity and (2) collection of suitable material
for future systematic research. Specimens were fixed
in 10% buffered formalin and later transferred to
alcohol. Color slides were taken of representative
live individuals of many species. Frog calls were
recorded during the 1992 survey. Collected material
was deposited in three research collections: the 1992
material at the Museum of Zoology, University of
Michigan (UMMZ), and Departement de Biologie
Animale, Universite d' Antananarivo (UADBA); and
the 1996 material at the American Museum of Nat-
ural History (AMNH) and UADBA.

Results

During our two surveys, a total of 5 1 amphib-
ian and 62 reptile species were recorded in the
PN de Marojejy, giving a total herpetofaunal di-

RASELIMANANA ET AL: HERPETOFAUNA

159

Table 9-1. Amphibians and reptiles identified during two surveys in five elevational zones within the PN de
Marojejy or at indicated sites at the periphery of the park.

Transect (m)

Elevation (m)

Species

450

750

1225 1625 1875

Min.

Max.

Amphibia
Microhylidae
Platypelis barbouri
Platypelis grandis
Platypelis occultans
Platypelis tsaratananaensis
Platypelis tuberifera
Plethodontohyla bipunctata
Plethodontohyla coudreaui
Plethodontohyla notosticta
Plethodontohyla ocellata
Plethodontohyla serratopalpebrosa
Stumpffia grandis
Stumpffia roseifemoralis
Stumpffia tridactyla
Stumpffia sp.

Hyperoliidae

Heterixalus madagascariensis
Ranidae

Ptychadena mascareniensis
Mantellidae

Laurentomantis malagasia

Mantella laevigata

Mantella nigricans

Mantella sp.

Mantidactylus aglavei

Mantidactylus albofrenatus

Mantidactylus asper

Mantidactylus betsileanus

Mantidactylus bicalcaratus

Mantidactylus biporus

Mantidactylus cornutus

Mantidactylus curtus

Mantidactylus femoralis

Mantidactylus grandidieri

Mantidactylus klemmeri

Mantidactylus leucomaculatus

Mantidactylus liber

Mantidactylus lugubris

Mantidactylus luteus

Mantidactylus opiparis

Mantidactylus peraccae

Mantidactylus pulcher

Mantidactylus redimitus

Mantidactylus rivicola

Mantidactylus ulcerosus

Mantidactylus sp.

Rhacophoridae
Boophis albilabris
Boophis anjanaharibeensis
Boophis brachychir
Boophis englaenderi
Boophis luteus
Boophis madagascariensis
Boophis marojezensis

Manantenina

Mandena

460

780

350

1200

350

1300

1560

1560

700

1200

300

550

770

1225

700

1300

470

470

1120

1870

1150

1300

680

1250

1550

1875

350

1225

80

80

70

100

3560

790

350

770

450

850

750

850

350

1150

300

1100

430

1225

720

800

710

1850

450

1550

680

1875

350

700

350

1900

100

500

430

1300

450

730

500

1550

450

460

360

780

450

450

1150

1675

750

1220

350

780

700

850

375

760

1875

1875

760

760

350

700

480

1550

450

650

450

780

350

740

500

1560

160

FIELDIANA: ZOOLOGY

Table 9- 1 . Continued.

Species

Transect (m)

450

750

1225 1625 1875

Elevation (m)

Min.

Max.

Boophis reticulatus
Boophis rappiodes
Boophis tephraeomystax
Boophis sp. 1
Boophis sp. 2

Reptilia
Gekkonidae
Ebenavia inunguis
Geckolepis maculata
Blaesodactylus antongilensis
Lygodactylus miops
Microscalabotes bivittis
Paroedura gracilis
Phelsuma guttata
Phelsuma laticauda
Phelsuma lineata
Phelsuma madagascariensis
Phelsuma pusilla
Phelsuma quadriocellata
Uroplatus ebenaui
Uroplatus fimbriatus
Uroplatus lineatus
Uroplatus sikorae
Uroplatus sp.

Chamaeleonidae
Brookesia betschi
Brookesia griveaudi
Brookesia karchei
Brookesia minima
Brookesia vadoni
Calumma cf. brevicornis
Calumma boettgeri
Calumma cucullata
Calumma gastrotaenia
Calumma cf. malthe
Calumma nasuta
Calumma peyrierasi
Furcifer pardalis

Scincidae
Amphiglossus astrolabi
Amphiglossus intermedius
Amphiglossus macrocercus
Amphiglossus mandokava
Amphiglossus melanopleura
Amphiglossus melanurus
Amphiglossus minutus
Amphiglossus mouroundavae
Amphiglossus ornaticeps
Amphiglossus punctatus
Mabuya gravenhorstii
Paracontias holomelas
Paracontias milloti
Pseudoacontias angelorum

Cordylidae
Zonosaurus madagascariensis
Zonosaurus rufipes
Zonosaurus subunicolor

Mandena

Mandena

1 km S of Mandena

Mandena

400

400

500

1560

80

80

350

350

525

525

350

730

350

375

350

520

470

750

480

480

350

800

250

850

100

100

1225

1850

150

375

100

600

710

800

1150

1950

360

750

350

720

720

1200

380

550

1150

1650

350

950

380

850

450

1550

850

1150

1800

2130

1100

1200

440

720

450

1675

1200

1570

350

1350

1675

1950

80

680

70

100

480

720

1225

1900

300

350

575

1220

470

700

450

1250

720

770

700

700

700

750

100

100

380

380

300

300

650

650

380

750

360

750

375

700

RASELIMANANA ET AL: HERPETOFAUNA

161

Table 9-1. Continued.

Species

Boidae

Boa manditra
Typhlopidae

Typhlops sp.

Typhlops ocularis

Colubridae
Alluaudina bellyi
Dromicodryas quadrilineatus
Geodipsas laphystia
Geodipsas sp.
Ithycyphus miniatus
Leioheterodon madagascariensis
Liophidium torquatus
Liophidium rhodogaster
Liopholidophis epistibes
Liopholidophis stumpffi
Liopholidophis sp.
Pararhadinaea albignaci
Pseudoxyrhopus heterurus
Pseudoxyrhopus microps
Pseudoxyrhopus tritaeniatus
Lycodryas arctifasciatus
Lycodryas betsileanus
Lycodryas gaimardi

Transect (m)

Elevation (m)

450

750

1225 1625 1875

Min.

Max.

350

780

2 km NW of Mandena 1

4 km E of Andapa

2 km NW of Mandena 1

1 km S of Mandena

470

700

640

700

150

380

200

200

350

790

470

1560

800

800

200

200

70

70

650

850

150

1600

350

600

1250

1875

300

300

350

350

700

790

750

750

740

740

350

350

700

740

Observation only.

versity of 113 species (Table 9-1). In addition, a
further three species of amphibians and seven spe-
cies of reptiles were recorded from outside but
within 5 km of the park boundary (Mandena,
Manantenina, or Andapa). Five amphibians and
five reptiles represented undescribed species. One
of these, Pseudoacontias angelorum, has been de-
scribed subsequently by Nussbaum and Raxwor-
thy (1995). We recorded 64 species that were not
previously known from the region, of which 57
species are new records for the park. This previ-
ously unknown component of the PN de Marojejy
herpetofauna represents 50% of the entire species
sampled by us and reflects both the high species
diversity of the area and the relatively low-inten-
sity surveying that has been conducted previously
within the park.

Maximum herpetofaunal species diversity was
recorded at mid-elevation, at the 750 m transect
(76 species), and minimum diversity was recorded
at the 1875 m transect (12 species; Fig. 9-1). A
total of 1,155 pitfall trap days yielded 39 individ-
ual amphibians and reptiles, giving an overall dai-
ly pitfall capture success rate of 3.4%. All 13 spe-
cies sampled by pitfalls were also captured by

other survey methods. The pitfall trap capture
data are given in Table 9-2.

Discussion

Species Recorded by Previous Surveys

Table 9-3 is a summary of the species previ-
ously reported from the PN de Marojejy in inven-
tory lists published by Nicoll and Langrand
(1989), Safford and Duckworth (1990), Blom-
mers-Schlosser and Blanc (1991), and Glaw and
Vences (1994, 1997). We could not confirm the
presence (inside the park) of 21 amphibian and 6
reptile species reported by these authors. Some of
these species are likely to have been misidentified,
such as Zonosaurus aeneus, or may reflect alter-
native taxonomic determinations within groups
(especially amphibians) that are easily confused.
Potential examples include Heterixalus betsileo,
Laurentomantis horrida, Mantidactylus majori,
M. pseudoasper, and M. spiniferus. Resolution of
these taxonomic problems will require examina-

162

FIELDIANA: ZOOLOGY

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RASELIMANANA ET AL: HERPETOFAUNA

163

40 n

(J)

O 20

I Amphibians
Reptiles

450 m 750 m

1225 m 1625 m 1875 m

Transect Zone
Fig. 9-1. Herpetofaunal diversity surveyed within each transect zone of the PN de Marojejy.

tion of the relevant collected material. In some
instances, frog calls and live photographs are also
required to confirm identifications, such as in the
Boophis goudoti group frogs. For example, we
suspect that some of our B. reticulatus material
from Marojejy contain B. rufioculis and B. bur-
geri specimens, but diagnostic characters for these
species include iris coloration and call character-
istics (Glaw & Vences, 1997), which are no lon-
ger evident in the collected material.

Some of the Marojejy records reported by
Blommers-Schlosser and Blanc (1991) and Glaw
and Vences (1994) may include material collected
outside the park. For example, we recorded eight
species (8% of the total herpetofauna sampled by
us in this area) at Mandena, Manantenina, or close
to Andapa that were not sampled within the park
itself. Finally, some Marojejy records may repre-
sent collecting efforts made on the eastern or
northern sides of the massif. Brygoo (1978) re-
corded Brookesia superciliaris at Marojejy on the
basis of a specimen from an unknown collector.
This chameleon was not found on the summit trail
during our surveys but was collected during a
brief 1994 visit to the northern slopes of the Ma-
rojejy Massif at Antsirandrano (Ramanamanjato,
pers. comm.). Because Brookesia superciliaris is
easy to sample at other sites, we suspect this cha-

meleon may be absent from the eastern part of the
PN de Marojejy, where we conducted our survey.

The herpetofaunal capture success rate of the
pitfall traps (3.4%) is comparable with other east-
ern rain forest sites such as Andringitra (3.5%),
Anjanaharibe-Sud (2.1%), and Andohahela
(3.0%) (Raxworthy & Nussbaum, 1996; Raxwor-
thy et al., 1998; Nussbaum et al., 1999). Trap suc-
cess rates decreased with increasing elevation:
1.8% for lines above 1500 m, compared with
4.2% for lines between 450 and 1250 m elevation.
No species was collected solely with pitfall traps,
unlike prior studies that have used identical pitfall
sampling methods (see references above). Proba-
bly the longer duration of transect surveying used
in this study increased the sampling success of the
other survey methods.

The species accumulation curves for each tran-
sect are plotted in Figure 9-2. For transects that
were surveyed during both 1992 and 1996, the
first part of the curve represents the 1992 sam-
pling period. The longest period of sampling was
28 days, for the 750 m transect (however, for 6
days between 27 November and 2 December
1992, the majority of survey effort was made at
other elevations), and the minimum period was 8
days for the 1875 m transect. The only transect
that reached an obvious plateau with regard to

164

FTELDIANA: ZOOLOGY

o

<D

a.

*5 45

o

r
12 14 16 1

Sampling Days

20

24

28

Fig. 9-2. Accumulation curves of herpetofaunal diversity (all sampling techniques) within each transect zone of
the PN de Marojejy. Arrows indicate the end of the 1992 survey and the start of the 1996 survey, for each transect.

newly accumulated species was the 1225 m tran-
sect, during the final 4 days of sampling. How-
ever, for all transects, species were being accu-
mulated at less than two species per day over the
final 2 to 6 days. Although these results indicate
that sampling cannot be considered complete for
any transect (with the possible exception of 1225
m), visual extrapolation of these accumulation
curves indicate that 80%-95% of the herpetofau-
na was sampled at each transect. The possibility
that we did not find up to 27 previously recorded
species within the park (but see comments above)
also suggests that our 75 total days of survey ef-
fort was not sufficient time to achieve a complete
herpetofaunal inventory of the PN de Marojejy.

Elevational Distribution

The large elevation range found within the
park makes this one of the best areas in Mada-
gascar to study elevational influences on pat-
terns of species distribution. The elevational ef-

fect on distribution is well illustrated by the
Chamaeleonidae. At elevations below 1000 m,
the following species occur: Brookesia griveau-
di, B. karchei, Calumma cucullata, and Furcifer
pardalis. Middle elevation specialists (850-
1650 m) are Brookesia betschi, B. vadoni, Ca-
lumma boettgeri, and C. cf. malthe. High-ele-
vation specialists (1650-2132 m) are Calumma
cf. brevicornis and C. peyrierasi. Three species
have broader elevational distributions, occur-
ring at both middle and lower elevations:
Brookesia minima, Calumma gastrotaenia, and
C. nasuta. Our surveys represent elevational
range extensions for some species, such as Ca-
lumma peyrierasi, previously recorded at 1900-
2000 m (Brygoo et al., 1974a) but found in this
study at 1675 m. Two subspecies of Calumma
gastrotaenia are known from within the park:
C. g. marojezensis at 600 m (Brygoo et al.,
1970a) and C. g. guillaumeti at 1300 m (Brygoo
et al., 1974a). We found Calumma gastrotaenia
marojezensis at the lower two transects, below
850 m, and C. g. guillaumeti at 1250-1675 m.
The allopatric elevational distribution of these

RASELIMANANA ET AL: HERPETOFAUNA

165

Table 9-3. Published inventory lists for the herpetofauna of the PN de Marojejy and massif.

Species

Blommers-

Glaw &

Nicoll &

Safford &

Schlosser

Vences

This studv

Langrand

Duckworth

& Blanc

(1994 &

(1992 &

(1989)

(1990)

(1991)

1997)

1996)

Amphibia
Microhylidae
Anodonthyla boulengeri
Dyscophus insularis
Platypelis barbouri
Platypelis grandis
Platypelis occultans
Platypelis pollicaris
Platypelis tsaratananensis
Platypelis tuberifera
Plethodontohyla bipunctata
Plethodontohyla coudreaui
Plethodontohyla minuta
Plethodontohyla notosticta
Plethodontohyla ocellata
Plethodontohyla serratopalpebrosa
Rhombophryne testudo
Stumpffia grandis
Stumpffia psologlossa
Stumpffia roseifemoralis
Stumpffia tetradactyla
Stumpffia tridactyla
Stumpffia sp.

Hyperoliidae
HeterLxalus betsileo
Heterixalus madagascariensis

Ranidae
Ptychadena mascareniensis

Mantellidae
Laurentomantis horrida
Laurentomantis malagasia
Mantella laevigata
Mantella madagascariensis
Mantella nigricans
Mantella sp.
Mantidactylus aglavei
Mantidactylus albofrenatus
Mantidactylus albolineatus
Mantidactylus asper
Mantidactylus betsileanus
Mantidactylus bicalcaratus
Mantidactylus biporus
Mantidactylus cornutus
Mantidactylus curtus
Mantidactylus femoralis
Mantidactylus flavobrunneus
Mantidactylus grandidieri
Mantidactylus grandisonae
Mantidactylus granulatus
Mantidactylus guttulatus
Mantidactylus klemineri
Mantidactylus leucomaculatus
Mantidactylus liber
Mantidactylus lugubris
Mantidactylus luteus
Mantidactylus opiparis

166

FLELDIANA: ZOOLOGY

Table 9-3. Continued.

Blommers-

Glaw &

Nicoll &

Safford &

Schlosser

Vences

This study

Langrand

Duckworth

& Blanc

(1994 &

(1992 &

Species

(1989)

(1990)

(1991)

1997)

1996)

Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus
Mantidactylus

majori

peraccae

pliciferus

pseudoasper

pulcher

punctatus

rivicola

redimitus

spiniferus

ulcerosus

sp.

Rhacophoridae
Boophis albilabris
Boophis anjanaharibeensis
Boophis brachychir
Boophis cf. burgeri
Boophis englaenderi
Boophis luteus
Boophis madagascariensis
Boophis majori
Boophis marojezensis
Boophis rappiodes
Boophis reticulatus
Boophis tephraeomystax
Boophis sp. 1
Boophis sp. 2

Reptilia
Gekkonidae
Ebenavia inunguis
Geckolepis maculata
Blaesodactylus antongilensis
Lygodactylus miops
Microscalabotes bivittis
Paroedura gracilis
Phelsuma guttata
Phelsuma laticauda
Phelsuma madagascariensis
Phelsuma lineata
Phelsuma pusilla
Phelsuma quadriocellata
Uroplatus ebenaui
Uroplatus fimbriatus
Uroplatus lineatus
Uroplatus sikorae
Uroplatus sp.

Chamaeleonidae
Brookesia betschi
Brookesia griveaudi
Brookesia karchei
Brookesia minima
Brookesia superciliaris
Brookesia vadoni
Calumma bifidus
Calumma cf. brevicornis
Calumma boettgeri
Calumma cucullata

RASELIMANANA ET AL: HERPETOFAUNA

167

Table 9-3. Continued.

Species

Nicoll &
Langrand

(1989)

Safford &

Duckworth

(1990)

Blommers-

Schlosser

& Blanc

(1991)

Glaw &

Vences

(1994 &

1997)

This study

(1992 &

1996)

Calumma gastrotaenia
Calumma globifer
Calumma cf. malthe
Calumma nasuta
Calumma peyrierasi
Furcifer pardalis

Scincidae
Amphiglossus astrolabi
Amphiglossus intermedins
Amphiglossus macrocercus
Amphiglossus mandokava
Amphiglossus melanopleura
Amphiglossus melanurus
Amphiglossus minutus
Amphiglossus mouroundavae
Amphiglossus ornaticeps
Amphiglossus punctatus
Mabuya gravenhorstii
Paracontias holomelas
Paracontias milloti
Pseudoacontias angelorum

Cordylidae
Zonosaurus aeneus
Zonosaurus madagascariensis
Zonosaurus rufipes
Zonosaurus subunicolor

Boidae

Boa manditra
Typhlopidae

Typhlops sp.

Typhlops ocularis

Colubridae
Alluaudina belly i
Dromicodryas quadrilineatus
Geodipsas infralineata
Geodipsas laphystia
Geodipsas sp.
Ithycyphus blanci
Ithycyphus miniatus
Leioheterodon madagascariensis
Liophidium torquatus
Liophidium rhodogaster
Liopholidophis epistibes
Liopholidophis stumpffi
Liopholidophis sp.
Pararhadinaea melanogaster
Pseudoxyrhopus heterurus
Pseudoxyrhopus microps
Pseudoxyrhopus tritaeniatus
Lycodryas arctifasciatus
Lycodryas betsileanus
Lycodryas gaimardi

*

*2

*
*2

Note The reported records in Nicoll and Langrand (1989) are based on published literature and personal obser-
vations. Those in Safford and Duckworth (1990) are based on personal observations. Those in Blommers-Schlosser

168

FIELDIANA: ZOOLOGY

~
o

CD
Q.
(J)

40n

35-

30-

25 -

Jr. 20-

15

10-

5-

I

wfrwiwiwi

Amphibians
Reptiles

3 4 5 6 7 8 9 101112131415161718192021

Elevation (100 m)
Fig. 9-3. Species diversity as a function of elevation, for amphibians and reptiles.

subspecies suggests that they are probably sibling
species, but the taxonomic status of these forms
awaits further study of the collected material.
Bohme et al. (1997) have recently suggested that
all Calumma gastrotaenia subspecies are indepen-
dent species.

Fig. 9-3 shows the influence of elevation on
species diversity. The species diversities of am-
phibians, reptiles, and both groups combined are
shown for 100 m elevational increments, with the
minimum and maximum elevations (recorded by
us in the PN de Marojejy) used to calculate the
elevational range of each species. This method
makes the assumption that each species is distrib-
uted continuously between the minimum and
maximum elevation recorded. The diversity of
amphibians and reptiles increases from the lower
elevation, peaking, for both groups, at the 700 m
elevational band (700-799 m). Above the 700 m
band, diversity drops off quickly until reduced to

just a single species {Calumma cf. brevicornis)
above 2000 m.

The 700 m Marojejy mid-elevation bulge in
species diversity is very similar to that described
by Raxworthy and Nussbaum (1996a) for the PN
d'Andringitra (maximum elevation 2658 m),
where maximum species diversity was recorded
at 800 m for amphibians and at 700 m for reptiles
(but, in this study, transects below 700 m were
not included, since they fell outside the park limit;
thus, the increase phase of the bulge was weakly
confirmed by only a single 100 m increment for
amphibians).

However, we note that the Marojejy mid-ele-
vation bulge in diversity might represent an arti-
fact of the greater sampling time spent at the 750
m elevation transect (28 days) than at the 450 and
1225 m transects (14 days). Despite the 14 extra
days of sampling at the 750 m transect, only three
more species were recorded than at the 450 m

and Blanc (1991) are based on museum specimens. Those in Glaw and Vences (1994) are based on published
literature, including Blommers-Schlosser and Blanc (1991), and additional collecting.

1 Authors almost certainly meant Stumpffia tridactyla (S. tetradactyla had not yet been described at that time).

2 Observations without vouchers, made by this survey.

RASELIMANANA ET AL: HERPETOFAUNA

169

Table 9-4. Distribution of Marojejy species (found in this study) at four other protected areas (see text for
literature sources).

Species

Montagne Anjanaharibe-

d'Ambre Sud Andringitra Andohahela

Amphibia
Microhylidae
Platypelis barbouri
Platypelis grandis
Platypelis occultans
Platypelis tsaratananaensis
Platypelis tuberifera
Plethodontohyla bipunctata
Plethodontohyla notosticta
Plethodontohyla serratopalpebrosa
Stumpffia grandis
Stumpffia roseifemoralis

Mantellidae

Mantidactylus aglavei
Mantidactylus albofrenatus
Mantidactylus asper
Mantidactylus betsileanus
Mantidactylus bicalcaratus
Mantidactylus biporus
Mantidactylus cornutus
Mantidactylus curtus
Mantidactylus femoralis
Mantidactylus grandidieri
Mantidactylus grandisonae
Mantidactylus klemmeri
Mantidactylus liber
Mantidactylus lugubris
Mantidactylus luteus
Mantidactylus opiparis
Mantidactylus peraccae
Mantidactylus pulcher
Mantidactylus redimitus
Mantidactylus rivicola
Mantidactylus ulcerosus

Rhacophoridae
Boophis albilabris
Boophis anjanaharibeensis
Boophis brachychir
Boophis luteus
Boophis madagascariensis
Boophis marojezensis
Boophis rappiodes
Boophis reticulatus

Reptilia
Gekkonidae
Ebenavia inunguis
Geckolepis maculata
Lygodactylus miops
Paroedura gracilis
Phelsuma guttata
Phelsuma lineata
Phelsuma madagascariensis
Phelsuma quadriocellata
Uroplatus ebenaui
Uroplatus fimbriatus
Uroplatus sikorae

170

FIELDIANA: ZOOLOGY

Table 9-4. Continued.

Species

Montagne Anjanaharibe-

d'Ambre Sud Andringitra Andohahela

Chamaeleonidae
Brookesia betschi
Brookesia vadoni
Calumma boettgeri
Calumma gastrotaenia
Calumma malthe
Calumma nasuta
Furcifer pardalis

Scincidae
Amphiglossus macrocercus
Amphiglossus melanopleura
Amphiglossus melanurus
Amphiglossus minutus
Amphiglossus mouroundavae
Amphiglossus ornaticeps
Amphiglossus punctatus
Mabuya gravenhorstii

Cordylidae
Zonosaurus madagascariensis

Boidae
Boa manditra

Colubridae
Alluaudina bellyi
Liophidium rhodogaster
Liopholidophis epistibes
Lycodryas arctifasciatus
Lycodryas betsileanus
Pseudoxyrhopus microps
Pseudoxyrhopus tritaeniatus

Total

26

*
61

37

28

transect (Fig. 9-1). For the first 14 days of sam-
pling, the species accumulation curves show
greatest species diversity at the 450 m transect
(Fig. 9-2). These sampling curves suggest species
diversity actually decreases with elevation for all
five of our surveyed transects. More data are
clearly needed to determine trends in herpeto-
faunal diversity below 800 m elevation, especially
to confirm the increase phase. Further discussion
of elevational patterns of species diversity is giv-
en by Raxworthy et al. (1998).

Comparisons with Other Sites

On the basis of our field surveys during the last
6 years, we provide here a comparison of the
shared herpetofaunal diversity between the PN de
Marojejy and four other protected rain forest ar-
eas: PN de la Montagne d'Ambre (Raxworthy &

Nussbaum, 1994); PN d'Andringitra (Raxworthy
& Nussbaum, 1996); Reserve Special (RS)
d'Anjanaharibe-Sud (Raxworthy et al., 1998); and
PN d' Andohahela (Nussbaum et al., 1999) (Table
9-4). The elevations surveyed at these sites varied
from 400 to 2300 m. These reserves share many
vegetation characteristics, such as moist rain for-
est at middle and low elevations and bamboo (and
in some cases ericoid heathland) at higher eleva-
tions. The majority of species shared among these
sites are low- and mid-elevation species. The PN
de Marojejy and the RS d'Anjanaharibe-Sud have
the highest number of shared species (61), and the
PN de Marojejy and the RNI d' Andohahela have
the lowest number (28). This corresponds to the
geographic distance between these sites. However,
the PN de Marojejy shares fewer species with the
PN de la Montagne d'Ambre, just 200 km from
Marojejy, than with PN d'Andringitra, which is
800 km from Marojejy. The herpetofauna of Mon-

RASELIMANANA ET AL: HERPETOFAUNA

171

tagne d'Ambre includes many species endemic to
northwestern Madagascar (sometimes called the
"Sambirano Domain"), whereas Andringitra and
Marojejy share many eastern endemic species. A
more detailed description of patterns of endemism
for the rain forest regions of Madagascar is given
by Rax worthy and Nussbaum (1996b, 1997).

The PN de Marojejy herpetofauna exhibit a sig-
nificant number of taxa that are possibly endemic
to the massif: Stumpffia sp., Mantella sp., Manti-
dactylus sp., Boophis sp. 1, Boophis sp. 2, Uro-
platus sp., Pseudoacontias angelorum, Brookesia
karchei, Calumma cf. brevicornis, C. gastrotaenia
guillaumeti, C. peyrierasi, Geodipsas sp., and
Liopholidophis sp. These 13 taxa make up 12%
of the park's herpetofauna, a degree of site en-
demism comparable with other sites, such as PN
d' Andringitra (13%; Raxworthy & Nussbaum,
1996a). Of additional significance was the discov-
ery of the rare and extremely poorly known gecko
Microscalabotes bivittis, which represents the first
record of this species in northern Madagascar (the
only other known localities are Perinet and the
Betsileo region; Pasteur, 1967).

Conservation Issues for the PN de
Marojejy Region

PN de Marojejy protects at least 113 species of
reptiles and amphibians. This is the highest di-
versity we have recorded so far within any of the
protected areas of Madagascar. It is also likely
that all 10 species that we found close to the park
boundary occur within this protected area and that
some of the 26 other species reported from Ma-
rojejy (that we failed to find) are also present. The
only protected area that is likely to have more
species than the PN de Marojejy is the RNI de
Tsaratanana, which has an even greater elevation-
al range (227 to 2876 m) and a comparable sur-
face area (48,622 ha) (Nicoll & Langrand, 1989).
These two "megadiversity" protected areas must
play a key role within the future protected areas
network of Madagascar.

The Andapa Basin, which is extremely favor-
able for paddy rice cultivation, has previously
provided some protection for the preservation of
forest on the western slopes of the park, by keep-
ing local cultivation activities centered on the ba-
sin itself. However, continuing demographic
growth, along with the migration of new settlers,
constitutes an increased threat because of the need

for new agricultural areas. For the eastern slopes
of the Marojejy Massif, slash and burn (tavy) is
an important agricultural method, and much of the
lowland forest in peripheral areas of the park has
now been transformed into agricultural areas. We
also noted many abandoned clearings produced by
slash-and-burn activities within the park, espe-
cially at elevations below 600 m. According to
one of our guides, this agricultural activity had
continued until 1994.

Hunting is another threat to the park. According
to our guides and some village elders, hunting
groups of 5 to 10 men live in the forest for at
least a week. These groups clear areas of forest
to make camping sites where they can build shel-
ters and dry bush meat. During our last visit, we
found many cleared areas and trails that were
made by hunters. The illicit exploitation of some
resources, such as lemurs, might not be sustain-
able, and the frequent passages of these hunters
into the park is also damaging the forest habitat
itself. As an example, Pandanus palm fronds are
used for the construction of temporary shelters,
but Pandanus also provides the microhabitat or
refugia for several species of amphibians and rep-
tiles within the PN de Marojejy, such as Manti-
dactylus pulcher, M. bicalcaratus, Platypelis tub-
erifera, Phelsuma lineata, and P. quadriocellata.

The recent change in status of Marojejy to a
Pare National will give local communities new
opportunities to generate income from ecotour-
ism, as tourists are allowed access to the park.
However, studies to assess the environmental im-
pact of ecotourism are urgently needed. The sum-
mit trail, currently the only access route to the
summit, will inevitably be used heavily by tourists
unless alternative routes are established. It is not
difficult to imagine this trail quickly becoming
damaged by tourists, making further research or
long-term monitoring impossible at the sites of
our transects. In view of the large investment of
scientific research that has already been made
along this trail over a period of 30 years, we
strongly recommend that an alternative tourist ac-
cess route be established within the newly created
park.

The PN de Marojejy is the main source of water
for many of the villages in the surrounding areas,
and this water source also offers the possibility
that irrigated paddy fields can be developed in the
deforested valley bottoms. Increased cultivation
of rice in paddy fields may help to reduce the
degree of slash-and-burn hill rice cultivation,
which continues to threaten the lowest-elevation

172

FTELDIANA: ZOOLOGY

forests at the periphery of the park. Similar to the
situation found in most of the rain forest protected
areas of Madagascar, the low-elevation herpeto-
fauna of the PN de Marojejy is the most vulner-
able to habitat loss or degradation; therefore, this
peripheral habitat of this park must be adequately
protected.

Acknowledgments

We thank the twins, Angelin Razafimanantsoa
and Angeluc Razafimanantsoa, Gabriella Rahari-
manana, and Bruno Zeze for their enthusiasm and
contribution to the field surveys. We also thank
Malalarisoa Razafimpahanana for help with edit-
ing the text, Steve Goodman and Jean-Paul Pad-
dack for constructive remarks on earlier versions
of the text, and Eleanor Sterling for coordinating
activities in New York. This field survey was pos-
sible because of the agreement of the Ministries
des Eaux et Forets and the Association Nationale
pour la Gestion des Aires Protegees, which kindly
delivered the requested authorizations. Research
support was provided to R. A. Nussbaum and C.
J. Raxworthy by the National Science Foundation
(DEB 90-24505, 93-22600) and the National Geo-
graphic Society (5396-94). The 1996 survey was
supported by the American Museum of Natural
History (Center for Biodiversity and Conserva-
tion) and the World Wide Fund for Nature.

Literature Cited

Bohme, W. 1997. Eine neue Chamaleonart aus der Cal-
umma gastrotaenia verwandtschaft Ost Madagas-
kars. Herpetofauna, 19: 5-10.

Blommers-Schlosser, R. M. A., and C. P. Blanc. 1991.
Amphibiens (premiere partie). Faune de Madagascar,
75(1): 1-385.

Brygoo, E. R. 1978. Reptiles Sauriens Chamaelonidae.
Genre Brookesia et complement pour le genre Cha-
maeleo. Faune de Madagascar, 47: 1-174.

Brygoo, E. R., C. P. Blanc, and C. A. Domergue.
1970a. Notes sur les Chamaeleo de Madagascar. VI.
C. gastrotaenia marojezensis n. subsp. d'un massif
montagneux du Nord-Est. Annales de l'Universite de
Madagascar, Serie Sciences de la Nature et Mathe-
matiques, 7: 273-278.

. 1970b. Notes sur les Brookesia de Madagascar.

III. B. karchei n.sp. Du massif du Marojezy. Annales
de l'Universite de Madagascar, Serie Sciences de la
Nature et Mathematiques, 7: 267-27 1 .

. 1974a. Notes sur les Chamaeleo de Madagascar

XII. Cameleons du Marojezy. C. peyrierasi n.sp. et C.
gastrotaenia guillaumeti n. subsp. (Reptilia, Squamata,
Chamaeleonidae). Bulletin de l'Academie Malgache,
51(1): 150-164.

1974b. Notes sur les Brookesia de Madagascar

VIII. Brookesia du Marojezy. B. betschi et B. gri-
veaudi n.sp. (Reptilia, Squamata, Chamaeleonidae).
Bulletin de l'Academie Malgache, 51(1): 151-166.
Domergue, C. A. 1984. Notes sur les serpents de la
region malgache. IV Le genre Pararhadinaea Boett-
ger, 1898. Desription d'une espece et d'une sous-es-
pece nouvelles. Bulletin du Museum National
d'Histoire Naturelle, Paris, 4th serie, sect. A, 6: 149
157.

1988. Notes sur les serpents de la region mal-

gache. VIII. Colubrides nouveaux. Bulletin du Mu-
seum National d'Histoire Naturelle, Paris, 4th serie,
sect. A, 10: 135-146.

Glaw, F, and M. Vences. 1994. A Fieldguide to the
Amphibians and Reptiles of Madagascar, 2nd ed.
Moss Druck, Leverkusen, Germany, 480 pp.

. 1997. Neue ergebnisse zur Boophis goudoti-

Gruppe aus Madagaskar: Bioakustik, fortpflanzungss-
trategien und beschreibung von Boophis rufioculis sp.
nov. Salamanandra, 32: 225-242.

Guillaumet, J.-L., J.-M. Betsch, C. Blanc, P. Morat,
and A. Peyrieras. 1975. Etude des ecosystemes mon-
tagnards dans la region malgache. III. Le Marojezy.
IV Lltremo et l'lbity. Geomorphologie, climatologie,
faune et flore (Campagne RCP 225, 1972-1973). Bul-
letin de Museum National d'Histoire Naturelle, 3rd
serie, 309(Ecologie Generate 25): 29-67.

Jenkins, M. 1987. Madagascar: An Environmental Pro-
file. IUCN, Cambridge, 374 pp.

Nicoll, M. E., and O. Langrand. 1989. Madagascar:
Revue de la Conservation et des Aires Protegees.
World Wide Fund for Nature, Gland, 374 pp.

Nussbaum, R. A., and C J. Raxworthy. 1995. Review
of the Scincine genus Pseudoacontias Barboza du Bo-
cage (Reptilia: Sauria: Scincidae) of Madagascar. Her-
petologica, 51(1): 91-99.

Nussbaum, R. A., C. J. Raxworthy, A. P. Raseliman-
ana, and J.-B. Ramanamanjato. 1999. Amphibians
and reptiles of the Reserve Naturelle Integrale
d'Andohahela, Madagascar, pp. 155-173. In Good-
man, S. M., ed., A Floral and Faunal Inventory of the
Reserve Naturelle Integrale d'Andohahela, Madagas-
car: With Reference to Elevational Variation. Fieldi-
ana: Zoology, n.s., 94: 1-297.

Pasteur, G. 1967. Redecouverte d'un genre de sauriens
malgache: Microscalabotes (Gekkonides). Annales de
l'Universite de Madagascar, Serie Sciences de la Na-
ture et Mathematiques, 5: 75-78.

Raxworthy, C. J., and R. A. Nussbaum. 1994. A rain-
forest survey of amphibians, reptiles and small mam-
mals at Montagne d'Ambre, Madagascar. Biological
Conservation, 69: 65-73.

. 1996a. Amphibians and reptiles of the Reserve

Naturelle Integrale d'Andringitra, Madagascar: A
study of elevational distribution and local endemicity,
pp. 158-170. In Goodman, S. M., ed., A Floral and
Faunal Inventory of the Eastern Slopes of the Reserve
Naturelle Integrale d'Andringitra, Madagascar: With

RASELIMANANA ET AL: HERPETOFAUNA

173

Reference to Elevational Variation. Fieldiana: Zoolo-
gy, n.s., 85: 1-319.

. 1996b. Patterns of endemism for terrestrial ver-

tebrates in eastern Madagascar, pp. 368-384. In Lour-
en^o, W. R., ed., Biogeographie de Madagascar. OR-
STOM, Paris.

. 1997. Biogeographie patterns of reptiles in east-
ern Madagascar, pp. 124-141. In Goodman, S. M..
and B. D. Patterson, eds, Natural Change and Human
Impact in Madagascar. Smithsonian Institution Press,
Washington, D.C.

Raxworthy, C. J., E Andreone, R. A. Nussbaum, N.
Rabibisoa, and H. Randriamahazo. 1998. Amphibi-
ans and reptiles of the Anjanaharibe Massif, Mada-
gascar: Elevational distribution and regional endemic-
ity, pp. 79-92. In Goodman, S. M., ed., A Floral and

Faunal Inventory of the Eastern Slopes of the Reserve
Speciale d'Anjanaharibe-Sud, Madagascar: With Ref-
erence to Elevational Variation. Fieldiana: Zoology,
n.s., 90: 1-246.

Safford, R., and W Duckworth. 1990. Systematic list
of reptiles recorded at Marojejy, pp. 136-138. In Saf-
ford, R., and W Duckworth, eds., A wildlife survey
of Marojejy Nature Reserve, Madagascar, Internation-
al Council for Bird Preservation study report no. 40.
International Council for Bird Preservation, Cam-
bridge, U.K.

Vences, M., E Glaw, and E Andreone. 1997. Descrip-
tions of two new frogs of the genus Mantidactylus
from Madagascar, with notes on Mantidactylus klem-
meri (Guibe, 1974) and Mantidactylus webbi (Gran-
dison, 1953) (Amphibia, Ranidae, Mantellinae). Aly-
tes, 14: 130-146.

174

FIELDIANA: ZOOLOGY

Chapter 10

Birds of the Pare National de Marojejy,
Madagascar: With Reference to Elevational
Distribution

Steven M. Goodman, 1 A. F. A. Hawkins, 2 and
Jean- Claude Razafimahaimodison 3

Abstract

Between 4 October and 20 November 1996, we evaluated bird species presence, relative
abundance, and bird community variation with altitude and vegetation type on the southeastern
slopes of the Pare National (PN) de Marojejy. This site, located in northeastern Madagascar,
occurs in the eastern humid formation and encompasses a wide variety of vegetational for-
mations. We collected data in elevational transect zones centered at 450, 775, 1250, 1625, and
1875 m, using mist-netting, point counts, call playback, and direct observation.

Eighty-four species, including 72 forest-limited birds, were recorded from the park. Species
richness decreased with increasing elevation, as did canopy height, whereas shrub density
increased. Of the 84 species recorded, 21 were detected only by direct observation and 1 only
by mist-netting. Call-playback recording was ineffective. The only new species recorded during
the survey for the reserve is Bernieria tenebrosa; Newtonia fanovanae was recorded subse-
quently. Mist-net capture rates in the PN de Marojejy were higher at the upper elevations than
at other sites. Arboreal frugivores were most abundant at lower elevations. In degraded forest
at 450 m, canopy and understory insectivores tended to be less frequent than in adjacent
primary forest. Survey efficiency may have been reduced by poor weather.

Resume

Entre le 4 octobre et le 20 novembre 1996, la presence des especes d'oiseaux, l'abondance
relative, et la variation de la communaute des oiseaux avec 1' altitude et le type de vegetation
ont ete evaluees sur les versants sud-est du Pare National de Marojejy. Ce site, situe au nord-
est de Madagascar, fait partie des formations humides de Test et contient une large variete de
ces formations vegetales. Les donnees sont collectees dans les zones de transects centrees a
450, 775, 1250, 1625, et 1875 m d'altitude, en utilisant les filets de capture, les comptages par
point, l'appel par play-back, et l'observation directe.

Quatre-vingt-quatre (84) especes dont 72 especes limitees aux forets ont ete recensees dans
le Pare. Sur ces 84 especes inventoriees, 21 ont ete decouvertes seulement par observation
directe et une espece par filet de capture. L'appel par play-back a ete inefficace. La seule
nouvelle especes recensee durant 1' inventaire a ete Bernieria tenebrosa; Newtonia fanovanae

1 Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, U.S.A.

2 BirdLife International, B.P. 1074, Antananarivo (101), Madagascar.

3 Department of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York,
NY 10024, U.S.A.

GOODMAN ET AL.: BIRDS 175

a ete trouve ulterieurement. Les seuils de captures par filet ont ete plus eleves dans les hautes
altitudes par rapport aux autres sites. Les frugivores arboricoles ont ete plus abondantes dans
les basses altitudes. Dans la foret degradee, a 450 m d'altitude, les insectivores des canopees
et des sous-bois tendaient a etre moins frequentes que dans la foret primaire voisine. Lefficacite
de l'inventaire pourrait etre reduite a cause du mauvais temps.

Introduction

In recent years there has been a rapid increase
in studies and available information on the extant
birds of Madagascar, including, for example, re-
discoveries of species that were thought to be ex-
ceptionally rare and perhaps on the verge of ex-
tinction (e.g., Thorstrom et al., 1995), descriptions
of two new species and a genus new to science
(Goodman et al., 1995, 1997a), studies on behav-
ior and breeding systems (Prum & Razafindratsita,
1997), and systematic studies (Schulenberg et al.,
1993; Fjeldsa et al., 1999; Goodman & Weight,
in press). Perhaps the best indication of this rapid
growth in information and general interest in the
birds of the island is the production of no less than
five field guides to Malagasy birds over the course
of the past decade (Langrand, 1990, 1995; Ya-
magishi et al., 1997; Morris & Hawkins, 1998;
Sinclair & Langrand, 1998). After primates, birds
are the most frequently researched group of ver-
tebrates in the protected areas system of Mada-
gascar (ANGAP, 1998). On the basis of the efforts
of national and foreign ornithologists working on
the island, the avifauna of Madagascar has be-
come much better understood over the past de-
cade.

Since the early 1990s, there has been a resur-
gence in interest among biologists in documenting
the fauna and flora of the island, particularly in
forested regions that have never been properly
surveyed or for groups that have not been ade-
quately studied. A number of recent inventories
have examined the distribution of biota along ele-
vational gradients on various mountains in the
eastern humid forests. All of these surveys were
conducted in protected areas or in soon-to-become
protected areas, both to document the plants and
animals occurring locally and to provide baseline
data for future biological monitoring. Ornitholog-
ical surveys were conducted at the following
sites (from north to south, Fig. 10-1): Pare Na-
tional (PN) de la Montagne d'Ambre (Goodman
et al., 1996a). the Reserve Speciale (RS)
d' Anjanaharibe-Sud (Hawkins et al., 1998), PN de
Masoala (Thorstrom & Watson, 1997), RS

d'Ambatovaky (Thompson & Evans, 1992), Re-
serve Naturelle Integrate (RNI) de Zahamena
(Hawkins et al., in press), PN d'Andringitra
(Goodman & Putnam, 1996; Goodman & Raso-
lonandrasana, in press), and the PN d'Andohahela
(Goodman et al., 1997b; Hawkins & Goodman,
1999). As part of this series of inventories, we
conducted a survey of the birds occurring in the
PN de Marojejy, the results of which are present-
ed here. Given that the sites listed above span the
complete length of the eastern humid forests of
the island, encompassing nearly 12 of latitude,
and given that the complete elevational range of
each mountain was surveyed, interesting ques-
tions can be addressed concerning the relationship
between altitudinal and latitudinal gradients over
a region that is relatively homogeneous with re-
gard to the botanical community (Koechlin et al.,
1974).

Review of Ornithological Research in
the Marojejy Region

The Marojejy Massif, one of the ornithologi-
cally best-known mountains on Madagascar, har-
bors a large number of endemic Malagasy bird
species. The current level of information on the
avifauna of the site is based on the work of nu-
merous researchers over the course of nearly 70
years. The following summary is a historical
overview of the ornithological research conducted
in the Marojejy region.

The earliest known avifaunistic work in the
Marojejy area was conducted by members of the
Franco-Anglo-Americaine (FAA) expedition, who
visited many different localities across the island
over the course of 26 months between 1929 and
1931. Between 19 August and 7 September 1930,
they collected specimens at a site "one day west"
of Andapa (Rand, 1932, 1936). This description
has been interpreted as indicating a locality lying
very close to the modern RS d' Anjanaharibe-Sud.
Although the camp was reported to be at 1800 m
and at the upper limit of rain forest, many of the

176

FIELDIANA: ZOOLOGY

46E

Tsaratanana ,
0,

14S

18S

22S

100 200 300

LM II I I J

km

50E

Marqjejy
Anjanaharibe-Sud

Masoala

Ambatovaky
Zahamena

Ranomafana

Andringitra

Andohahela

wm Eastern Humid Forest

I I Western Deciduous Forest

1 1 Southern Spiny Forest

Fig. 10-1. Map showing the position of the Pare National de Marojejy and several other sites in the eastern humid
forest in which the bird faunas have been studied along elevational transects.

species they reported from the site are character-
istic, on the basis of specimen material, of lower
elevations. It is probable that specimens were
brought in from a wide range of altitudes (Haw-
kins et al., 1998). The bird specimens collected
by the Mission FAA were divided between the
American Museum of Natural History, New York;
the British Museum (Natural History) now
known as the Natural History Museum, London;

and the Museum National d'Histoire Naturelle,
Paris.

In 1958, P. Griveaud visited the reserve for en-
tomological studies, but also made some obser-
vations on the avifauna (Griveaud, 1960). He also
made a small collection of birds. We have been
unable to locate the Griveaud collection in a nat-
ural history museum. In September 1972, an or-
nithological team visited the southeastern portion

GOODMAN ET AL.: BIRDS

177

of the reserve and made several important obser-
vations, which are reported in Benson et al. (1976,
1977).

On the basis of a 1987 visit to the Marojejy
Massif, made in the context of the World Wide
Fund for Nature-Madagascar biodiversity and
protected areas program, Nicoll and Langrand
(1989) reported a list of 103 bird species. This list
was partially reproduced by Jenkins (1990).

Between August and late October 1988, a stu-
dent expedition from Cambridge University (Unit-
ed Kingdom), in collaboration with Malagasy re-
searchers, conducted a biological inventory of
several different sites within the reserve (Safford
& Duckworth, 1990). They reported the presence
of 104 bird species in and around the reserve. One
of the many noteworthy findings of this group was
the rediscovery of the Madagascar Serpent Eagle,
Euthorchis astur, which had not been reliably re-
corded since the 1930s (Sheldon & Duckworth,
1990).

Between mid-October and early December
1994, a multidisciplinary expedition conducted a
biological inventory of the RS d'Anjanaharibe-
Sud, about 40 km from the western slopes of the
PN de Marojejy. This group visited four different
sites on the eastern slopes of the Anjanaharibe-
Sud Massif, between 875 m and the summital
zone at 1950 m. The ornithological results of this
survey have been published as two chapters in a
monograph on the site: a general overview with
natural history information and density estimates
for selected species (Hawkins et al., 1998), and
information on the birds of prey (Thiollay, 1998).
The ornithological data from the RS
d'Anjanaharibe-Sud provide an important oppor-
tunity for comparison with data obtained in the
PN de Marojejy, and cross-references between
these two sites are made throughout the current
chapter.

Goals of the Study

In light of the intense rate of deforestation in
Madagascar, it is critical to understand aspects of
the island's bird fauna in relation to habitat re-
quirements, geographical and elevational distri-
bution, and relative densities. Our study had four
main research objectives: (1) to add to previously
available information on the birds occurring in the
PN de Marojejy; (2) to determine the composition
of bird communities across elevational gradients;

(3) to determine the relative abundance of forest
species at different elevations; and (4) to compare
various surveyed mountains on the eastern portion
of the island with respect to the relationship be-
tween elevation and species richness along lati-
tudinal gradients.

Study Site and Methods

Study Site

Our survey of the PN de Marojejy was con-
ducted as part of the multidisciplinary study of
the biota of the mountain, between 4 October and
20 November 1996. (For an overview of the sur-
vey and for more precise botanical details of the
various study sites, see Chapters 1 and 3.) Orni-
thological data were collected within an altitudi-
nal interval of 75 m elevation, centered around
transect zones at 450, 775, 1250, 1625, and 1875
m. These transect zones were placed within 3 km
horizontal walking distance from our camps 1
(450 m), 2 (775 m), 3 (1325 m), 4 (1550 m), and
5 (1875 m). Throughout this chapter, we refer to
the elevational zones with respect to the place-
ment of the camps.

Most of the camp sites were situated near rivers
or streams in closed-canopy forest. The lowest
camp (450 m) was between two small rivers that
are tributaries of the Manantenina River. The veg-
etation type at this site was largely dominated by
indigenous native lowland forest, although exten-
sive areas of secondary forest were present. At
the 775 m site, which was at a site called Andam-
pimbazaha on an open rocky outcrop along the
Ambavaomby River, the forest showed few signs
of human disturbance and contained more floristic
elements of montane forest than did the 450 m
site (see Chapter 3). A marked change in the veg-
etational structure occurred between the 775 and
1250 m zones. Clear montane elements were pre-
sent at this latter site: the canopy height was low-
er, epiphytic loads were heavier, and ground
mosses were abundant. By 1625 m the forest was
completely dominated by montane elements, in-
cluding some sclerophyllous plants, and the forest
was further reduced in stature. Tree root systems
often formed hollow cavities under a superficial
cover of mosses and lichens, especially along the
ridges and slopes. The 1875 m site encompassed
the ecotone between the upper limit of sclero-
phyllous forest and open grassland savanna. The

178

FIELDIANA: ZOOLOGY

Table 10-1. Details of ornithological surveys in
each elevational zone within the PN de Marojejy.

Dura-

Species

Eleva-

tion

res-

tional

of

Species

species

tric-

zone

visit
(days)

Point
counts

record-
ed

ted to

(m)

No. (%)

zone

450 m

10

19

64

59 (92.2)

7

775 m

10

22

53

51 (96.2)

2

1250 m

10

22

57

55 (96.5)

1

1625 m

9

16

40

37 (92.5)

1875 m

7

11

32

22 (68.8)

6

Table 10-2. Accumulation of bird species previous-
ly unrecorded within each elevational zone in the PN de
Marojejy.

Eleva-
tional

zone 1

Cumulative no. of species
recorded by working day

8

10

450 m 13 40 43 55 57 61 64 64 64 64

775 m 9 20 41 46 53 53 53 53 53 53

1250 m 4 13 33 44 45 57 57 57 57 57

1625 m 18 20 28 35 36 39 40 40 40

1875 m 14 28 32 32 32 32 32 32

camp site at 1 875 m was on a ridge above the tree
line and slightly above a small forest-lined stream
(Andranomifototra). Within the open area, there
were large exposed outcrops of rock, some cov-
ered by small geophytes. More detailed botanical
descriptions of each transect zone are provided in
Chapter 3.

Methods

Survey methods used to collect information
about the birds of the PN de Marojejy included
direct observation, point counts, static observa-
tions from areas with broken canopy cover, mist-
netting, and call playback. To maximize infor-
mation from each zone, particularly for the gen-
eral observations, point counts, and mist-netting
procedures, different microtopographical and mi-
crohabitat areas were surveyed (e.g., ridges,
slopes, and valley bottoms).

J.-C. Razafimahaimodison was the principal
field ornithologist on the mission. A. F. A. Haw-
kins was present for the 450 and 775 m samples
and, together with Razafimahaimodison, con-
ducted point counts and general observations.
In higher altitude zones, Razafimahaimodison
was solely responsible for the point counts. In
all transect zones, S. M. Goodman, in collabo-
ration with M. Ravokatra, was responsible for
the bird netting and secondarily for bird obser-
vations. During inclement weather, such as
strong wind and heavy rain, point counts were
not conducted.

Survey Methods

Point Counts Point counts were used to es-
timate relative bird species frequency (Reynolds

et al., 1980; Bibby et al., 1992) occurring within
each elevational zone surveyed. Sample points
were located at 150 m intervals along preexisting
or recently cut trails. We conducted a minimum
of 15 point counts within each elevational zone
except the 1875 m zone, in which only 11 point-
count sites were installed, because of the limited
area of forest.

Each point-count site was sampled twice on
different days, once between 0500 and 0630 hr
and once between 0630 and 0900 hr, for a period
of 10 minutes. A 1 -minute "rest period" was used
for the equilibration of bird activity after arrival
at a station (Reynolds et al., 1980). All bird con-
tacts, visual and aural, were noted, and the hori-
zontal distance to the observer was estimated in
intervals of 0-5, 5-10, 10-20, 20-30, 30-50, and
50-100 m; distances greater than 100 m were not
precisely estimated. In cases of unfamiliar bird-
song, a tape recorder with a directional micro-
phone was used to record the vocalization for later
identification. If necessary, after a point-count
session was finished, recorded calls and vocali-
zations of unfamiliar birds were played back, to
attract the birds and thereby enable visual deter-
mination of the species involved.

Direct Observation We made direct obser-
vations while walking along preestablished trails
or bushwhacking in the forest. We noted bird spe-
cies, nature of contact (auditory or visual), type
of vocalization (song or call), the number of in-
dividuals seen or heard, behavior, foods con-
sumed, and other information. In addition, mixed-
species bird flocks were followed. Active search-
ing was conducted after the daily point counts. At
least 7 hours per day were spent in active searches
for birds.

Canopy Observations Additional observa-
tions were made from spots with broken-canopy
vistas to identify raptors and other bird species

GOODMAN ET AL.: BIRDS

179

Table 10-3. Elevational distribution of birds recorded in the elevational transects in the PN de Marojejy during
the 1996 survey. 1 with information on their distributional status 2 and habitat use. 3

Presence in elevational zone

Species

450 m

775 m 1250 m 1625 m 1875 m

Outside
reserve

Tachybaptus ruficollis
Anhinga rufa@
Ardeola idae*
Ardeola ralloides
Egretta alba@
Egret ta dimorpha@
Ardea purpurea
9cLophotibis cristata*
9cAviceda madagascariensis*
9cPolyboroides radiatus*
9cAccipiter henstii*
9cAccipiter francesii( * )
9cButeo brachypterus*
Falco newtoni(*)
Falco peregrinus@
Falco eleonoraeft
Margaroperdix madagascariensis*
Coturnix coturnix
9cMesitornis unicolor*
9cCanirallus kioloides*
9cSarothrura insularis*
9cAlectroenas madagascariensis*
9cStreptopelia picturata(*)
9cTreron australis{*)
Agapornis cana*
9cCoracopsis spp. 4 (*)
9cCoua caerulea*
9cCoua reynaudii*
9cCoua serriana*
9cCoua cristata*
9cCuculus rochii*
9cCentropus toulou(*)
9cOtus rutilusi*)
9cNinox superciliaris*
9cAsio madagascariensis*
Caprimulgus madagascariensis*
9cZoonavena grandidieri(*)
Apus melba@
Apus barbatus@
Cypsiurus panus
Merops superciliosus
Alcedo vintsioides{*)
9dspidina madagascariensis*
9cEurystomus glaucurus*
9cLeptosomus discolor^*)
9cBrachypteracias leptosomus*
9cBrachypteracias squamiger*
9cAtelornis pittoides*
9cAtelornis crossleyi*
9cPhilepitta castanea*
9cNeodrepanis coruscans*
9cNeodrepanis hypoxantha*
Mirafra hova*
Phedina borbonica(*)
9cMotacilla flaviventris*
9cCoracina cinerea*
9cHypsipetes madagascariensis(*)

+

+

+

-

+

+

_

+

-

+

-

+

(+)
(+)
(+)

(+)

180

FIELDIANA: ZOOLOGY

Table 10-3. Continued.

Species

Presence at elevational zone

450 m

775 m

1250 m 1625 m

1875 m

Outside
reserve

%Bernieria cinereiceps*
%Bernieria zoster ops*
%Bernieria tenebrosa*
%Bernieria madagascariensis*
%Bernieria xanthophrys*
%Calicalicus madagascariensis*
%Vanga curvirostris*
%Leptopterus viridis*
%Leptopterus chabert*
%Cyanolanius madagascarinus{*)
%Oriolia bernieri*
%Euryceros prevostii*
%Tylas eduardi*
Saxicola torquata
%Copsychus albospecularis*
%Monticola sharpei*
%Neomixis striatigula*
%Neomixis viridis*
%Neomixis tenella*
Cisticola cherinai*)
%Cryptosylvicola randrianasoloi*
%Oxylabes madagascariensis*
%Mystacornis crossleyi*
Acrocephalus newtoni*
%Nesillas typica(*)
%Dromaeocercus brunneus*
%Randia pseudozosterops*
%Newtonia brunneicauda*
%Newtonia amphichroa*
%Pseudobias wardi*
%Terpsiphone mutata(*)
%Nectarinia notata(*)
%Nectarinia souimanga{*)
%Zosterops maderaspatana(*)
Lonchura nana*
%Ploceus nelicourvi*
%Foudia madagascariensis*
%Foudia omissa*
%Dicrurus forficatus(*)
%Hartlaubius auratus*

Total no. of species
Total no. of forest species

-

-

+

+

+

+
+
+

+

-

-

+

+

-

-

+

+

-

+

+

+

+

(+)

+

+

+

-

-

+

+

+

+

(+)

+

+

-

-

-

+
+
+

-

+

-

-

+

+

+

+

+

+

+

(+)

-

-

-

+

+

+

+

+

-

-

-

-

+

+

(+)

+

+

+

+

-

+

+

+

+

-

+

+

+

-

-

-

+

+

+

+

+

+

+

-

+

+

+

+

-

+

+

+

+

+

-

-

+

+

+

+

+

+

+

-

+

+

+

+

+

+

+

+

+

+

+

+

-

+

-

+

+

+

+

-

+

+

+

-

-

+

+

+

+

+

+
+
+

+

+

+

+

+

+

+

+

+

-

+

+

+

+

-

-

+

+

+

+

64

53

57

40

32

59

51

55

37

22

Note. Records are also presented for species observed in areas just outside the reserve limits.

1 Key to presence/absence status: + = species recorded in zone; - = species not recorded in zone. Records from
the 1875 m zone in parentheses are of species recorded only in the lower portion of the transect, below tree line.

2 Key to distributional status: * = breeding species endemic to Madagascar; (*) = breeding species endemic to
Madagascar and nearby islands; # = migrant to Madagascar; @ = breeding species also occurring outside of region.

3 Key to habitat use: % = forest-dependent species (species depends on forest for a portion of its life cycle).

4 Because of difficulties in distinguishing between Coracopsis vasa and C. nigra in the field, we have combined
these species here. The vast majority of our records concern C. nigra.

that fly above the trees. Such sites were found
along rivers, on top of rocky outcrops, on ridges,
and in other open areas. We spent at least 10 hours
within each elevational zone conducting this type
of census.

Mist-Netting Mist-netting was used to pro-
vide information on the abundance of individual
species, as expressed by the number of captures
per net-day (Karr, 1981). Ten nets were set up at
each of the five elevational zones and were run

GOODMAN ET AL.: BIRDS

181

continuously for 5 days. Sites were generally
within 500-750 m trail distance from the camps.
All nets were 36 mm mesh size, 2.6 m high, and
12 m long. In most cases they were left open dur-
ing the night, to capture nocturnal bird species
and bats. They were checked nearly every hour
from sunrise to sunset and occasionally during the
night.

Most captured birds were weighed, measured,
and marked before being released near the origi-
nal site of capture. A mark was made with indel-
ible ink on the bird's primary feathers. Within
each elevational zone, the first primary feather
was marked for birds captured on the first day of
netting, the second primary feather on the second
day, and so on, until the fifth and final day of
netting. This marking technique provided a means
to recognize recaptured individuals and indicated
the chronology of the recaptures. Some birds were
collected and prepared as standard study skins,
skeletons, or fluid-preserved specimens. Tissue
samples of collected individuals were preserved
in a solution of EDTA. The specimens are depos-
ited in the Field Museum of Natural History, Chi-
cago, and the Departement de Biologie Animale,
Universite d' Antananarivo, Antananarivo.

Vegetation Structure Measurement

At each point-count site, the following vegeta-
tion structure measurements were made: (1) hab-
itat type (ridge, slope, or valley) and condition of
forest (primary, degraded, secondary, or grass-
land); (2) estimated canopy height and estimated
percentage canopy cover; and (3) shrub layer den-
sity, estimated by taking the mean of four esti-
mates of distance visible through the shrub layer
at eye level.

Systematic Order, Nomenclature, and
Common Names

We have generally followed the systematic or-
der, scientific names, and vernacular names of
Langrand (1995). The major exceptions include a
generic revision of Pseudocossyphus, now placed
in the genus Monticola (Goodman & Weight, in
press), and that of Phyllastrephus and Crossleyia,
now placed in the genus Bernieria (Fjeldsa et al.,
1999). For the latter two genera, to reflect this
taxonomic change, we resurrect vernacular names
proposed by earlier authors.

Results

For each of the five transect zones, information
on the duration of each survey, the number of
point-count sites, and number of species recorded
at each site is presented in Table 10-1. At most
of the sites, 5-6 days was sufficient to reach an
apparent plateau in the species discovery curve
(Table 10-2). The major exception was the 1625
m zone, where inclement weather reduced the rate
of species accumulation. We often spent the first
and second days in each elevational zone instal-
ling the camp, opening trails, and choosing sites
for the point counts.

Overall, 84 species of birds were recorded dur-
ing the 1996 inventory of the PN de Marojejy, 72
(86%) of which are forest-dwelling forms and 60
(71%) of which are endemic to Madagascar. An
additional 18 (22%) species breed in the Mala-
gasy region, 5 (6%) nest on the island but have
much broader distributions, and 1 (1%) is a Hol-
arctic migrant (Table 10-3). Thirteen species were
recorded in the area just outside the reserve,
which gives a total of 97 species recorded during
the 1996 inventory.

Several species of birds previously reported
from the PN de Marojejy were not recorded dur-
ing the 1996 inventory. These comprise a variety
of non-forest-dwelling species, including six
open-country, four aquatic, and eight predomi-
nantly forest-dwelling species (Table 10-4). When
this tally is added to the list produced during the
1 996 survey, the known avifauna of the park and
immediate surroundings includes 115 species.

The elevational distributions of bird species ob-
served during this inventory are presented in Ta-
ble 10-3. In total, 64 species were recorded in the
450 m zone, 53 in the 775 m zone, 57 in the 1250
m zone, 40 in the 1625 m zone, and 32 in the
1875 m zone. This pattern of decreasing bird spe-
cies richness with increasing elevation is typical
of eastern humid forests on the island (Hawkins,
1999).

The vast majority of species recorded during
the inventory had broad elevational distributions
across the Marojejy Massif. In a few cases, certain
species were noted only in a single elevational
zone (Table 10-3). Two species were restricted to
the 450 m zone {Treron austral is and Oriolia ber-
nieri). Both species tend to be lowland forest spe-
cies, and this result is concordant with their
known altitudinal distribution (Hawkins, 1999).
Further, Lonchura nana is not a forest species,
and its occurrence in this zone is almost certainly

182

FIELDIANA: ZOOLOGY

Table 10-4. Species not recorded during 1996 sur-
vey of the PN de Marojejy, but previously reported from

Table 10-5. Vegetation structure measures at point-
count sample sites in the PN de Marojejy.

Mean

Mean

Mean

Previ-

Altitude and

canopy

canopy

shrub

ous

vegetation

height

cover

density 1

re-

category

(m)

(%)

(m)

Species

Habitat

cord

450 m

Tachybaptus pelzelnii

Aquatic

1

Degraded forest

Bubulcus ibis
Butorides striatus
Milvus migrans
Accipiter madagascariensis

Open areas
Aquatic
Open areas
Forest

1
1

1,2
1

Ridge (N =
Valley (N =
Slope (N =

3)
5)
3)

18.3
18.0
11.0

56
36

5

6.0
4.8
3.1

Eutriorchis astur

Forest

2

Nondegraded forest

Falco concolor

Open areas

1

Ridge (N =

3)

31.5

80

7.6

Falco zoniventris

Forest, open

1,2

Valley (N -

2)

27.5

80

5.8

areas

Slope (N =

3)

30.0

90

8.0

Turnix nigricollis

Open areas

1

775 m

Rallus madagascariensis

Aquatic

1,3

Dryolimnas cuvieri

Aquatic

1,2

Ridge (N =

4)

23.0

90

5.5

Caprimulgus enarratus

Forest

1,2

Valley (N =

6)

25.0

80

6.9

Riparia paludicola

Open areas

1

Slope (N =

12)

20.0

80

5.1

Hartertula flavoviridis

Forest

1,2

1250 m

Schetba rufa
Hypositta corallirostris
Xenopirostris polleni
Corvus albus

Forest
Forest
Forest
Open areas

1,2
1,2
4
1,2

Ridge (N =
Valley (N =
Slope (N =

1625 m

9)

5)
8)

10.0
20.0
15.0

60
80
90

3.0
6.0

5.3

Source of previous records:

1 = Nicoll and Langrand

Ridge (N =

3)

5.0

25

1.9

(1989); 2 = Safford and Duckworth (1990); 3

= Gri-

Valley (N =

4)

15.0

75

5.3

veaud (1960); 4 Benson et al. (1977).

Slope (N =

9)

10.6

60

3.7

1875 m

related to the open areas

associated with

forest

(all) (N

6)

4.0

40

3.1

degradation. Six other species (Polyboroides ra-
diatus, Accipiter francesii, Treron australis, Falco
peregrinus, Alcedo vintsiodes, and Hartlaubius
auratus) generally have much broader altitudinal
distributions on other surveyed mountains on
Madagascar (Hawkins, 1999), and we assume the
same holds for the Marojejy Massif. Two species
that tend to be uncommon were restricted to the
775 m zone: Mesitornis unicolor is known to oc-
cur from lowland to middle elevations in other
areas of the eastern humid forest, but it was re-
corded only at 860 m in the nearby RS
d'Anjanaharibe-Sud (Hawkins et al., 1998), and
Bernieria tenebrosa tends to be a lowland species
(Langrand, 1995). The single species restricted to
the 1 250 m zone, Eurystomus glaucurus, normally
occurs across a broad elevational zone (Langrand,
1995). Six species were recorded only in the 1875
m zone (Falco eleonorae, Margaroperdix mada-
gascariensis, Coturnix coturnix, Sarothrura insu-
laris, Caprimulgus madagascariensis, and Miraf-
ra hova), none of which are forest-restricted birds,
and these species occurred on the Marojejy Mas-
sif in the zone near or above forest line. In the
cases of the galliforms, nightjar, and lark, this is

1 Estimated by taking the mean of four estimates of
distance visible through the shrub layer at eye level (see
p. 182).

almost certainly related to the available habitat,
whereas the case of the falcon is probably due to
the broad, open expanses, in which such birds are
much easier to observe than in closed forest. Nor-
mally the rail has a broad elevational distribution
on the island, from near sea level to 2300 m (Lan-
grand, 1995), and we can offer no clear explana-
tion as to why it should be limited to the upper
portions of the massif. It was not recorded during
the 1988 expedition to the site (Safford & Duck-
worth, 1990), but it appears on Griveaud's list of
Marojejy species. Madagascar lacks species of
birds restricted to the zone above tree line as
found in other areas of the world with high moun-
tains, such as the Andes, Himalayas, and sites on
the African continent (Maclean, 1990; Inskipp &
Inskipp, 1991; Stotz et al., 1996). This is largely
because the highest peak on the island is slightly
more than 2800 m, the surface area of habitat
above tree line is rather restricted, and, in recent

GOODMAN ET AL.: BIRDS

183

Table 10-6. Netting results by elevational transect in the PN de Marojejy.

No. of individuals netted by

altitude

Species

450 m

775 m

1250 m

1625 m

1875 m

Total

Accipiter francesii

1

1

Otus rutilus

2

1

3

Alcedo vintsioides

1

1

Ispidina madagascariensis

2

4

6

Brachypteracias leptosomus

1

1

Philepitta castanea

3

16

11

2

32

Neodrepanis coruscans

4

9

13

Neodrepanis hypoxantha

10

1

11

Bernieria madagascariensis

4

4

Bernieria zosterops

2

2

4

Bernieria cinereiceps

7

7

14

Bernieria xanthophrys

2

1

3

Hypsipetes madagascariensis

2

1

3

Copsychus albospecularis

1

10

11

Monticola sharpei

3

2

5

Saxicola torquata

5

5

Nesillas typica

9

4

8

21

Newtonia amphichroa

2

7

5

14

Newtonia brunneicauda

2

2

Cryptosylvicola randrianasoloi

2

2

Terpsiphone mutata

3

4

5

12

Oxylabes madagascariensis

7

7

Nectarinia souimanga

5

5

4

3

5

22

Zosterops maderaspatana

6

2

2

1

2

13

Vanga curvirostris

3

3

Euryceros prevostii

1

1

Ploceus nelicoun'i

1

1

1

3

Foudia madagascariensis

7

7

Foudia omissa

4

3

1

8

Total no. of individuals

34

71

72

26

26

232

Total no. of species

13

16

16

9

6

28

Average no. of birds/net-day

0.7

1.4

1.4

0.5

0.6

0.9

Note. Netting results are for 50 cumulative net-days at each zone, for a total of 250 net-days. Figures do not
include recaptures.

geological periods, there have been considerable
shifts in the elevation of montane habitats as a
function of warm and cool periods (Straka, 1996).

Important vegetational differences were record-
ed in the different zones in which the transects
were conducted (see Chapter 3 for more details).
Mean canopy cover and height declined with in-
creasing elevation, and, between 1250 m and
1625 m, the canopy height tended to be highest
in valley bottoms, moderate on slopes, and lowest
on ridges (Table 10-5). Shrub density increased
with increasing elevation and was thickest on
ridges in the 1250 and 1625 m samples. There
was a dramatic difference between degraded and
intact forest in the 450 m sample; the degraded
forest had a lower and more open canopy and a
much more dense shrub layer.

In each elevational zone, 50 net-days were ac-

crued (Table 10-6). We captured a total of 232
individual birds belonging to 28 species. The
numbers of individuals and species captured were
highest at the 775 and 1 250 m sites, slightly lower
at 450 m, and notably lower at 1625 and 1875 m.
The weight data for captured birds are presented
in Table 10-7.

Discussion

Methodological Considerations

We recorded 84 bird species in the survey area
of the PN de Marojejy during the 1996 expedi-
tion, using three different methods: point counts
(59 species [70%]), direct observation (about 75

184

FTELDIANA: ZOOLOGY

Table 10-7. Mass (g) of birds netted during the survey of the PN de Marojejy.

Species

N

Mass (g), Range'

Mean SD 2

Accipiter f ranee sii

1

166(9)

Otus rutilus

1

114.5

Alcedo vintsioides

1

16.5

Ispidina madagascariensis

5

16.0-19.5

18.1 1.4

Philepitta castanea

31

24.5-43.5

35.3 3.7

Neodrepanis coruscans

10

4.5-7.5

6.5 1.0

Neodrepanis hypoxantha

11

6.5-9.0

7.0 0.7

Bernieria madagascariensis

4

22.0-34.5

27.9 6.5

Bernieria zosterops

3

19.5, 20.5,

21.0

Bernieria cinereiceps

13

13.5-20.0

18.3 1.7

Bernieria xanthophrys

3

17.0, 18.5,

20.5

Hypsipetes madagascariensis

3

41.5,41.5,

41.5

Copsychus albospecularis

11

20.0-28.5

24.7 2.3

Monticola sharpei

5

23.0-27.0

24.9 1.8

Saxicola torquata

5

12.0-14.5

13.4 1.1

Nesillas typica

20

14.0-22.0

18.3 2.4

Newtonia amphichroa

14

9.0-15.0

11.8 1.7

Newtonia brunneicauda

2

9.5, 13.5

Cryptosylvicola randrianasoloi

2

7.1,7.4

Terpsiphone mutata

12

10.0-15.5

13.8 1.5

Oxylabes madagascariensis

7

21.0-26.5

23.1 2.3

Nectarinia souimanga

22

6.5-10.5

7.6 1.0

Zosterops maderaspatana

13

9.0-13.0

10.5 1.0

Vanga curvirostris

3

58.5, 65.0,

70.0

Euryceros prevostii

1

87

Ploceus nelicourvi

3

22.5, 26.0,

30.0

When three or fewer measurements were available, only the masses themselves are given.
SD = standard deviation.

species [89%]), and mist-netting (28 species
[33%]). Among these 84 species, 21 (25%) were
recorded only from direct observation and 1 spe-
cies (1%) only from mist-netting. The results from
the three methods used during the field expedition
(point count, direct observation, and mist-netting)
are therefore complementary. Of the three tech-
niques, mist-netting provided the least amount of
total or unique information on the presence of
birds within any elevational transect. For this
technique to be more effective, a far greater num-
ber of nets must be run in each elevational zone.
Further, several important biases are inherent in
use of this technique to measure relative abun-
dance of birds (Remsen & Good, 1996).

The use of playback recordings of bird vocali-
zations provided mixed results during this survey.
For example, it was not possible to attract Mesi-
tornis unicolor with this technique, although it
was used numerous times in an area of forest
where this species was known to occur. However,
with some other species, such as Bernieria tene-
brosa, the recordings attracted the birds, and we
were able to confirm the identifications when they
came closer. Ispidina madagascariensis was the

only bird recorded by mist-netting but not by the
other techniques. This species is a secretive, un-
derstory animal that is easily overlooked.

The bird species recorded by general observa-
tions but not during point-count observations can
be divided into four groups: generally rare or un-
common species that were observed on only a few
occasions: Lophotibis cristata, Mesitornis unicol-
or, and Bernieria tenebrosa; high aerial foraging
species that were difficult to see in the dense for-
est in which most point-count sites were located:
Phedina borbonica, Apus barbatus, and Cypsi-
urus parvus; nocturnal species that were not ac-
tive during the period in which point counts were
conducted: Asio madagascariensis, Ninox super-
ciliaris, and Caprimulgus madagascariensis; and
forest raptors that are easily overlooked, particu-
larly when they occur in relatively low densities
and fly over the forest canopy: Aviceda madagas-
cariensis and Polyboroides radiatus.

Calculation of Point-Count Frequencies

The method used for the point counts was in-
tended to provide sufficient data for the calcula-

GOODMAN ET AL.: BIRDS

185

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GOODMAN ET AL.: BIRDS

187

tion of densities. However, as can be seen from
Table 10-8, contact frequencies with all but the
most common species were low. Densities cal-
culated on the basis of fewer than 20 contacts are
subject to very large error estimates, making com-
parisons with other data (e.g., for ecological mon-
itoring) unreliable. In this study, the only species
for which density estimates would be usable for
comparative purposes is Nectarinia souimanga,
an abundant species in degraded and secondary
habitats as well as in forest and, thus, of little use
as an indicator. A possible reason for the low level
of contacts may have been the unseasonably cold
and wet weather we experienced at the first four
camps.

To permit crude relative abundance compari-
sons between species and sites, we have calculat-
ed a contact frequency per sample for each spe-
cies detected on point counts (Table 10-8). This
figure is simply the overall number of contacts per
elevational sample divided by the number of
point-count sites within that sample.

Comparisons with Other Rain Forest
Inventories

Eighty-four species were recorded during the
1996 expedition to the PN de Marojejy, whereas
during the 1988 inventory, 104 species were ob-
served in and around the reserve (Evans et al.,
1992). The only forest species newly identified in
the reserve during the 1996 survey was Bernieria
tenebrosa. Sarothrura insularis was not recorded
by the 1988 group, but it had been noted previ-
ously on the massif (Griveaud, 1960; Nicoll &
Langrand, 1989). A number of species are noted
in the literature from the reserve (Griveaud, 1960;
Benson et al., 1976, 1977; Nicoll & Langrand,
1989; Evans et al., 1992) that were not recorded
during the 1996 inventory. Approximately half of
these birds are open-country or aquatic forms (Ta-
ble 10-4). Newtonia fanovanae was recorded by
Hawkins in 1997 in the western portion of the
reserve. The only species recorded in and around
the nearby RS d'Anjanaharibe-Sud that has not
yet been reported from the PN de Marojejy is Tyto
soumagnei (Hawkins et al., 1998).

For the most part, capture rates for the mist-
netting operations in the RS d'Anjanaharibe-Sud
and PN de Marojejy were comparable within sim-
ilar elevational zones. At the former site, the num-
ber of accrued net-days was 45 per elevational
zone, and at the latter site, 50 per elevational

zone. On the Anjanaharibe-Sud Massif, at 875 m,
the average number of individuals captured per
net-day was 1.0 (46 individuals of 13 species); on
the Marojejy Massif, at 775 m, 1.4 individuals per
net-day (71 individuals of 16 species). At 1260
m, 1 .0 individuals were captured per net-day (46
individuals of 13 species) and, at 1250 m, 1.4 in-
dividuals per net-day (72 individuals of 16 spe-
cies), respectively; at 1550 m, 1.9 individuals per
net-day (85 individuals of 15 species) and, at
1625 m, 0.5 individuals (9 species) per net-day,
respectively; at 1950 m, 0.6 individuals per net-
day (27 individuals of 8 species) and, at 1875 m,
0.6 individuals per net-day (29 individuals of six
species), respectively. Thus, the major differences
between the massifs seem to be in the montane
forest, where the capture rate in the RS
d'Anjanaharibe-Sud (1550 m) was more than
three times greater than in the PN de Marojejy
(1625 m). Factors such as differences in season
and associated variation in bird density or differ-
ences in canopy height between the two sites can-
not be invoked to explain the variation between
these two sites.

When all available information on the birds
documented in and around the PN de Marojejy
(Tables 10-3 and 10-4) is combined, the total
number of recorded forest species is 79. To our
knowledge, this is the most diverse forest bird
species list for any single mountain site on Mad-
agascar. In comparison with other massifs on
which elevational transects have been conducted,
69 forest bird species were recorded between 720
and 2450 m in the PN d'Andringitra (Goodman
& Rasolonandrasana, in press), 74 between 875
and 1950 m in the RS d'Anjanaharibe-Sud (Haw-
kins et al., 1998), 69 between 440 and 1875 m in
parcel 1 of the PN d'Andohahela (Goodman et al.,
1997b), and 73 between 450 and 1500 m in the
RNI de Zahamena (Hawkins et al., in press).

Although the elevational range surveyed for
birds on each of these mountains is not the same
and the amount of effort at each of these sites is
not really comparable, some generalizations can
be offered. The three sites that are perhaps the
most similar with regard to elevational range and
work effort are (from north to south) the PN de
Marojejy, the RNI d'Andringitra, and the PN
d'Andohahela (parcel 1), in which 79, 69, and 69
species of birds have been recorded, respectively.
On the basis of these results and other inferences,
it is clear that the humid forest portions of north-
ern Madagascar are ornithologically richer than
the southern portion of the island.

FIELDIANA: ZOOLOGY

70

60

50

40

30-

20

10-

D Montagnc d'Ambre

O Anjanaharibe-Sud

---O Marojcjy

- A Zahamena

- -ffl Andringitra

* Andohahela

400

600

800

1000 1200

1400

1600 1800 2000 2200 2400 2600

Elevation (m)

Fig. 10-2. Comparison of bird species richness as a function of elevation at several mountainous sites on Mad-
agascar. Data are from Montagne d'Ambre (Goodman et al., 1996), Anjanaharibe-Sud (Hawkins et al., 1998), Marojejy
(herein, Nicoll & Langrand, 1989; Evans et al., 1992), Zahamena (Hawkins et al., in press), Andringitra (Goodman
& Rasolonandrasana, in press), and Andohahela (Goodman et al., 1997b).

A series of plots of each of these sites with
regard to the number of bird species recorded per
elevational zone indicates a consistent pattern of
decreasing species richness with increasing ele-
vation (Figure 10-2). Deviations from this pattern
include the PN de la Montagne d'Ambre, in
which there was a greater number of species at
1000 m than at 340 m. However, this shift largely
reflects a change in forest type the lowland for-
ests of this mountain are distinctly drier than the
montane zone (Goodman et al., 1996). Evidence
of a slight mid-elevational bulge has been record-
ed in the PN d' Andohahela (parcel 1) at 810 m
and in the PN de Marojejy at 1250 m.

The Frugivorous Bird Community

The greatest number of large frugivorous bird
species was found in the lowland habitat at 450
m, where all three species of columbids known
from the Marojejy Massif (Alectroenas madagas-
cariensis, Streptopelia picturata, and Treron aus-
tralis) were found, as well as both Coracopsis
parrots. Treron, a species that appears to feed ex-
tensively on fruits, particularly those of Ficus, and
perhaps the most obligate large frugivore in the
eastern humid forest, did not occur above 450 m.
The highest altitude at which Streptopelia and
Alectroenas were recorded during the 1996 sur-

vey was 1250 m, although these species are
known to occur at higher elevations elsewhere on
Madagascar (Hawkins, 1999). The diet of the for-
mer species is composed of a high proportion of
seeds (Langrand, 1995; Goodman & Parrillo,
1997), and, probably for this reason, it has a
broader elevational distribution on the massif.
Further, Alectroenas is known to make large-scale
movements, and within forest habitat they are not
sedentary. The Coracopsis parrots often consume
seeds, flowers, and other food resources besides
fruits. The other species of frugivore on the Ma-
rojejy Massif, Philepitta castanea, which feeds on
small understory fruits (Goodman & Putnam,
1996; Razafindratsita, 1995), occurred across a
broad elevational range, from 450 to 1625 m.

On the basis of line transects and general col-
lections made by N. Messmer and P. J. Rakoto-
malaza (Chapter 3), the 450 m site had the largest
number of Ficus trees in fruit of any of the zones
during the period of the inventory. However, the
highest density of these trees was in the 775 m
zone, and this genus was not found at elevations
above 1200 m. On the basis of this data and the
importance of Ficus in the diet of Treron, we sus-
pect that, as the season advanced and Ficus fruit-
ed at higher zones, this pigeon would become
more common at higher elevations.

In summary, for large frugivores such as Tre-
ron or for other species that often feed exten-

GOODMAN ET AL.: BIRDS

189

sively on fruits, such as Alectroenas, Streptopelia,
and Coracopsis spp. particular life-history traits
need to be invoked to explain their elevational
distribution on the Marojejy Massif as well as on
other mountains in the eastern humid forest. The
two reasons for this are closely linked: (1) few
obligate frugivores occur in this forest type on
Madagascar, compared with other areas of tropical
forest, and (2) the phenology and distribution of
fruiting trees does not provide a consistent pattern
of fruit availability to allow the evolution of sed-
entary obligate frugivores (Goodman & Gan-
zhorn, 1997). Thus, more specifically, Treron
consumes proportionately more fruit and is lim-
ited to lower altitudes, where this food source,
particularly Moraceae, is more abundant, whereas
the other large semifrugivorous species (Alec-
troenas, Streptopelia, and Coracopsis spp.) have
a broader diet preference and accordingly occur
across a wider elevational zone, perhaps with sea-
sonal altitudinal migrations based on the avail-
ability of fruit resources.

Congeneric Overlap and Replacement

The altitudinal distribution of the two sunbird-
asities, Neodrepanis coruscans and N. hypoxan-
tha, appear to abut rather than overlap in the 1250
m zone. Parallel cases for the replacement of these
two species have been found on several moun-
tains where they occur along the same versant;
irrespective of latitude, the zone is generally be-
tween 1200 and 1300 m (e.g., Goodman & Put-
nam, 1996; Goodman et al., 1997b). At some sites
the transition zone occurs at higher elevations:
at 1500-1600 m on the slopes of the RS
d'Anjanaharibe-Sud (Hawkins et al., 1998) and at
1 800 m in the RNI de Tsaratanana (Hawkins, un-
publ. data). It would be interesting to study what
factors are responsible for this consistent and
abrupt shift. Perhaps this is the zone of transition
of important food plants, such as Bakerella (Lor-
anthaceae).

Three species of ground-rollers were recorded
in the 1250 m zone; this is discussed below, in
the species accounts, under Brachypteracias lep-
tosomus (p. 192).

The most speciose genus of forest-dwelling
birds in the PN de Marojejy is Bernieria (formerly
consisting of the genera Phyllastrephus and Cros-
sleyia), composed of five species. Two of these
birds, B. xanthophrys and B. tenebrosa, were rare
on the slopes of Marojejy, and we hesitate to infer

too much on the basis of limited information. The
former species appears to be a bird of montane
forests, occurring between approximately 1000
and 2000 m (Hawkins, 1999). During the 1996
survey, it was recorded in the 1250 and 1625 m
zones. The latter species is a lowland species, not-
ed only in the 775 m zone. The balance of the
three members of this genus have been previously
noted to show signs of both syntopy and eleva-
tional replacement on other mountains on Mada-
gascar (e.g., Goodman & Putnam, 1996; Good-
man et al., 1997b; Hawkins et al., 1998). The gen-
eral pattern is for B. madagascariensis and B. zos-
terops to occur together in lowland habitats and
to be largely or completely replaced by B. ciner-
eiceps at elevations above 1200 or 1300 m. This
pattern has been documented at sites in the north
from the Marojejy and Anjanaharibe-Sud Massifs
south to the PN d' Andohahela. Although this zone
covers nearly 10 of latitude, the altitude of re-
placement is relatively consistent between these
sites. In the PN d' Andohahela, the three species
occurred sympatrically from 810 to 1200 m.

Another interesting case is that of the genus
Neomixis. At all of the sites surveyed to date in
the eastern humid forest, where N. striatigula, N.
viridis, and N. tenella occur along the same
slopes, a relatively consistent pattern emerges,
once again irrespective of latitudinal gradients. N.
striatigula and N. viridis are found across a broad
elevational gradient, from lowland forest to at
least montane forest and sometimes to forest line
(Goodman et al., 1997b; Goodman & Rasolon-
andrasana, in press). N. tenella is a bird of low-
land to middle elevation forest, to approximately
1200 m and, rarely, to 1500 m.

Bird Communities in Intact and Degraded
Forest

In the 450 m zone, point-count sample sites
were approximately equally divided between rel-
atively intact forest and degraded forest, which
were intimately interdigitated on a scale of hun-
dreds of meters. The latter habitat was largely
composed of a mixture of bamboo and Aframo-
mum sp., with a few relict canopy trees. Evidently
this area was burned for cultivation before the es-
tablishment of the reserve in 1952. The fact that
degraded and intact forest were present so close
together permitted a comparison of the bird com-
munities of each without the usual problems of
controlling for larger-scale variability in other

190

FIELDIANA: ZOOLOGY

ecological parameters, such as soil type, original
vegetation type, exposure, and altitude.

In Table 10-8 we show the frequency of con-
tacts of species on point-count samples in degrad-
ed and intact forest. The number of species re-
corded in each habitat is not markedly different
(29 in degraded and 34 in intact forest), but there
is a marked difference in the distribution of cer-
tain species. Four species (Streptopelia picturata,
Ploceus nelicourvi, Oxylabes madagascariensis,
and Philepitta castanea) were not recorded on
point counts in degraded forest but were recorded
once each in intact forest. Meanwhile, Lonchura
nana and Leptopterus chabert were recorded once
each in the degraded sector and not in the intact
sector. The significance of these records is diffi-
cult to judge, since contact frequencies in these
samples are too low to allow much extrapolation
from these data. It is certainly true that Oxylabes
and Philepitta can be thought of as forest interior
species and that Lonchura and Leptopterus are
most common outside forest. Although differenc-
es in frequencies of other species between de-
graded and intact forest are sometimes large, they
are insufficient for statistical testing. However, we
can draw inferences from these disparities that
merit further investigation.

Nine other species were recorded more than
twice as often in one or the other habitat. Cuculus
rochii, Coua caerulea, Brachypteracias leptoso-
mus, Bernieria zosterops, Newtonia brunneicau-
da, Calicalicus madagascariensis, Leptopterus
viridis, and Euryceros prevostii all were much
less frequent or absent entirely from the degraded
area. Several of these taxa are generally either
canopy insectivores (Cuculus, Newtonia, Calical-
icus, and Leptopterus), which may be unable to
find sufficient food resources in degraded forest
with an open canopy (Table 10-5), or understory
insectivores (Coua, Brachypteracias, Bernieria,
and Euryceros), which depend on finding prey in
rather open primary forest understory and not in
the dense vegetation found in the degraded areas
(Table 10-5). It is striking that no understory in-
sectivore was recorded in the degraded forest.

Only one species, Randia pseudozosterops, was
recorded more than twice as frequently in degrad-
ed forest; all individuals detected in degraded for-
est were singing from the tops of isolated trees in
areas of bamboo or Aframomum. The significance
of this finding is difficult to evaluate. This species
is absent from degraded areas far from primary
forest (Morris & Hawkins, 1998), and it may only
be the presence of primary forest less than 100 m

from the degraded forest in which these birds
were singing that permitted them to use the de-
graded area.

Another point about the ability of certain birds
to colonize open areas is demonstrated by some
of the species occurring in the zone above tree
line. Mirafra hova is common to abundant in her-
baceous savanna habitats. It was not recorded in
any of the degraded open areas in the 450 m tran-
sect or in any other forested area in the reserve,
but it was relatively common from tree line to the
summital zone of the Marojejy Massif. The case
of Foudia madagascariensis is slightly different.
This species is common in open habitats across
most of the island and seems able to penetrate
forest areas through either natural or human-re-
lated habitat disturbance. Within forest habitat, F.
madagascariensis tends to occur along river mar-
gins and at the ecotone between open areas and
forest. It was recorded in the three lowest zones,
was absent in the 1 625 m zone, and was abundant
in the open area above forest line. In both cases,
these two species of birds are able to colonize
open habitats above forest line, although their ele-
vational distribution is not continuous across the
mountain. Thus, they are clearly dispersing across
habitats in which they do not otherwise occur.

Species Accounts

In this section we provide details on particular
bird species of interest. These include those clas-
sified as threatened or near threatened by Collar
et al. (1994). One aspect of the growing amount
of information on the birds of Madagascar is that
many species previously designated as rare are
now known to be relatively common but restricted
to particular habitats or elevational zones. This
point is discussed in more detail in the Conser-
vation Considerations section at the end of this
chapter.

Madagascar Crested Ibis, Lophotibis cristata

The Madagascar Crested Ibis was recorded on
a few occasions in the 450 and 775 m zones. It
was recorded at 1 300 m in this same sector of the
PN de Marojejy in 1988 (Safford & Duckworth,
1990). The density of this species in the reserve
was low; it is unclear whether this reflects natural
levels or is related to habitat disturbance and hunt-

GOODMAN ET AL.: BIRDS

191

ing pressure in the lower-lying areas of forest.
Few observations of L. cristata were made during
the inventory of the RS d'Anjanaharibe-Sud
(Hawkins et al., 1998).

Madagascar Cuckoo-falcon, Aviceda
madagascariensis

This species was observed on a few occasions
in the 1250, 1625, and 1875 m zones and near the
summit, at 1975 m. In the literature, the maximum
elevation of this species was previously noted as
1800 m (Rand, 1936). This raptor usually fre-
quents open areas and forest edge. In the RS
d'Anjanaharibe-Sud, it was recorded between 875
and 1550 m (Hawkins et al., 1998).

Madagascar Serpent-eagle, Eutriorchis astur

Until recently, this species was thought to be
very rare and was classified as threatened with
extinction (Sheldon & Duckworth, 1990). More
recently, after more than 50 years of not being
documented in the wild, it has been found at sev-
eral forested sites in the northern portion of the
island, including the RS d'Ambatovaky (Raxwor-
thy & Colston. 1992; Thompson & Evans, 1992).
the Marojejy forest (Sheldon & Duckworth.
1990), the RS d'Anjanaharibe-Sud (Thiollay.
1998), the Masoala Peninsula (Thorstrom et al.,
1995). and the RNI de Zahamena (Hawkins et al.,
in press).

During our 1996 expedition to the reserve, an
exhaustive search in different parts of the forest
was conducted, but no sign of this species was
discovered. This effort included many hours of
canopy searching from the Andampimbazaha out-
crop (the site of the 775 m camp), a site with a
panoramic view of birds flying over the canopy.
The fact that this species was not recorded in the
reserve during the 1996 expedition, although it
has been previously recorded in the region, prob-
ably indicates that densities are extremely low or
that this species is difficult to detect. Several re-
cent records of this species are based on birds
discovered vocalizing. The inclement weather ex-
perienced at the lowland camps during our survey
of the Marojejy Massif may have reduced the call-
ing frequency of this species and hence its de-
tectability.

Henst's Goshawk, Accipiter henstii

Henst's Goshawk was recorded rarely during
the survey. Records from our inventory include a
single bird at 450 m and one individual heard vo-
calizing at 1625 m for approximately 15 minutes.
In general, almost all contacts with this species
involve calling birds, and it is possible that the
inclement weather during the mission reduced vo-
calizations. Safford and Duckworth (1990) re-
ported fairly regular contact with this species in
the same sector of the reserve. In the nearby RS
d'Anjanaharibe-Sud, it was observed on a few oc-
casions at elevations between 875 and 1950 m
(Hawkins et al., 1998).

Brown Mesite, Mesitornis unicolor

This forest-dependent species was recorded
only in the 775 m zone, between the elevational
range of 725 and 875 m. On 16 October, it was
observed and heard calling in a forested valley
with dense undergrowth. During the next few
days, playback song of this species was broadcast
with a tape recorder to attract the bird, but it did
not respond. During the 1988 survey of Marojejy.
this species seemed equally uncommon (Safford
& Duckworth. 1990). In the RS d'Anjanaharibe-
Sud. this species was recorded only in the 875 m
zone (Hawkins et al.. 1998).

Short-legged Ground-roller, Brachypteracias
leptosomus

Records of this species from the Marojejy for-
est were obtained at the four lowest sites surveyed
and encompassed an elevational range from 450
to 1625 m. In the 450 m zone, one individual was
observed singing approximately 5 m from the
camp site. Two individuals were heard calling
each morning at approximately 0430 hr, not far
from the camp site.

Playback song was also used to attract this spe-
cies. In the 1250 m zone, one individual ap-
proached to within 2 m of the observer and re-
mained perched 1 m above the ground for ap-
proximately 30 minutes. It stopped calling for a
period and then resumed. At the same time this
bird was vocalizing, another individual of this
species was heard calling nearby. The Pitta-like
Ground-roller (Atelornis pirtoides) was also ob-
served in this same area of forest. Finally, a third

192

FIELDIANA: ZOOLOGY

species of ground-roller, Crossley's Ground-roller
(A. crossleyi), was found in this same zone. Thus,
three species of ground-roller occur sympatrically
within this elevational stratum. At other sites in
the eastern humid forest, the same three species
have been recorded in sympatry, including the
1260 m zone of the RS d'Anjanaharibe-Sud
(Hawkins et al., 1998) and the 1200 m zone of
the PN d'Andohahela (Goodman et al., 1997b).

Scaly Ground-roller, Brachypteracias
squamiger

This species was recorded in the Marojejy for-
est between 450 and 775 m. It is generally con-
sidered a bird of lowland forest, below 1000 m
(Langrand, 1995). A nest site of the Scaly
Ground-roller was found at approximately 450 m,
in an area of slightly degraded forest with steep
slopes and rolling hills. The nest was a hole, ap-
proximately 10 cm in diameter, approximately 1
m above ground level in a nearly vertical earth
bank in a shallow valley. On 1 1 October, an adult
was seen and heard calling as it approached this
nest. This species is broadly distributed across
intact zones of lowland forest, from the Marojejy
and Anjanaharibe-Sud Massifs south to PN
d'Andohahela (Goodman et al., 1997b; Hawkins
et al., 1998).

Pitta-like Ground-roller, Atelornis pittoides

In general, this species is the most common of
the four ground-rollers occurring in the humid
forests of eastern Madagascar. In the PN de Ma-
rojejy, it was rarely noted during our 1996 survey.
Records were obtained in the 1250 and 1625 m
zones. During the 1988 inventory of the reserve,
this species was regularly recorded in the south-
western and northwestern sectors but was not re-
corded in the southeastern sector (Safford &
Duckworth, 1990), which includes the area of our
1996 survey. It was also recorded only once in the
RS d'Anjanaharibe-Sud during 2 months of field
work in 1994. Further south for example, in the
PN de Ranomafana and PN d'Andohahela it is
distinctly more common (J. C. Razafimahaimo-
dison, pers. obs.; Goodman et al., 1997b; Hawkins
& Goodman, 1999). This species is often more
common than other species of ground-rollers in
drier and more degraded forests.

Rufous-headed Ground-roller, Atelornis
crossleyi

This ground-roller was recorded between 1250
and 1875 m. In the 1250 m zone, this species was
observed in a variety of microhabitats, including
ridges, valleys, and, less frequently, on slopes.
This species was regularly heard calling in the
early morning, often starting at 0400 hr. In the
1625 m transect, it appeared not to be uncommon
and was observed almost daily.

Common Sunbird-asity, Neodrepanis
coruscans

This species was recorded from 450 to 1250 m
and, on the basis of point-count data, appears to
be more common toward the upper limit of this
elevational range. (See next account for more de-
tails on this species' elevational range.)

Yellow-bellied Sunbird-asity, Neodrepanis
hypoxantha

The Yellow-bellied Sunbird-asity was recorded
within the elevational swath from 1125 to ap-
proximately 1775 m. The highest recorded obser-
vation occurred just below the ecotone between
the sclerophyllous forest and the open savanna.
Five individuals were caught in mist nets: four at
1250 m and one at 1625 m. The contact frequency
of this species in the former zone was much high-
er than in the latter (Table 10-8). Although both
species of Neodrepanis were observed within the
1250 m transect, they appear to have abutting ele-
vational distributions. N. coruscans was found
only in the lower end of the 1 250 m zone, down
to lowland forest, and N. hypoxantha was found
only in the upper portion of the 1250 m zone, up
to forest line. During the 1988 mission to this re-
serve, N. hypoxantha was observed on six occa-
sions between 1500 and 1700 m (Safford & Duck-
worth, 1990). Further, in the RS d'Anjanaharibe-
Sud, this species was recorded between 1500 and
1950 m (Hawkins et al., 1998). Thus, in the PN
de Marojejy, the number of observations and net-
ted individuals from the 1250 m zone seems
slightly lower than previously noted for this spe-
cies in northern Madagascar.

GOODMAN ET AL.: BIRDS

193

Brown Emu-tail, Dromaeocercus brunneus

The Brown Emu-tail was recorded in the higher
elevational zones of the PN de Marojejy, from
1125 to 1800 m. Generally we had only fleeting
glimpses of this terrestrial species as it scurried
away in dense vegetation. One individual was
captured in a live trap set for small mammals in
the ecotone between sclerophyllous forest and the
open grassland zone. The entrance of the trap was
embedded in a tunnel passageway in the ground,
and the only way for the bird to have entered the
trap was as it was exiting the burrow. During the
1988 Cambridge survey of the Marojejy forest,
this species was relatively common in the same
zone of forest we worked, with up to 10 observa-
tions per day between 1300 and 2000 m (Safford
& Duckworth, 1990). In the RS d'Anjanaharibe-
Sud, D. brunneus is broadly distributed, from 875
to 1950 m, being distinctly more common at high-
er elevations (Hawkins et al., 1998).

Rand's Warbler, R undid pseudozosterops

forest of the Andohahela Massif (Goodman &
Schulenberg, 1991), and in the northeast, in the
RS d'Ambatovaky (Evans, 1991). Since then it
has been found (from north to south) in the RS
d'Anjanaharibe-Sud (Hawkins et al., 1998), in the
RNI de Zahamena (Hawkins et al., in press), and
in the region in and around the PN d' Andohahela
(Goodman et al., 1997b).

Neither the 1988 nor the 1996 inventories of
the PN de Marojejy detected this species. In July
1997, A. F. A. Hawkins saw an individual Red-
tailed Newtonia in a mixed-species flock at ap-
proximately 500 m, in the far western portion of
the reserve, on the trail from Ambalamanasy to
Doany. This site is in the corridor linking the Ma-
rojejy Forest with the large forest block to the
west, which includes the RS d'Anjanaharibe-Sud.
Despite its loud and characteristic song, this spe-
cies has not been located in numerous other ap-
parently suitable eastern humid forest localities.
From the above data, it appears to be most com-
mon at low elevations, with patchy distribution
within this habitat.

In the Marojejy forest, Rand's Warbler was re-
corded within the altitudinal zone from 450 to
1625 m. This species always frequented the can-
opy zone of the forest.

Cryptic Warbler, Cryptosylvicola
randrianasoloi

Ward's Flycatcher, Pseudobias wardi

This species was recorded in the altitudinal
swath between 325 and 1675 m. It was generally
observed flying short distances between trees, in
dense forests along the riverside or at forest edge.

In the PN de Marojejy, from 1250 to 1875 m,
the Cryptic Warbler was a common member of
the local bird community. In the latter zone, this
species occurred in the highest forested corridors
hugging stream lines and surrounded by open
grassland. It was noted numerous times complete-
ly out of the forest zone, perched on top of grass
tufts and singing. The lower limit of this species
is above the 775 m zone and below the 1250 m
zone, which corroborates its elevational distribu-
tion on other mountains on the island (Goodman
et al., 1995, 1997b; Hawkins et al., 1998).

Red-tailed Newtonia, Newtonia fanovanae

This species was recently known from only a
single specimen captured in 1931 in the Sihanaka
Forest (in the region between Didy and Fito). In
1989, it was rediscovered in the southeast, in the

Dusky Tetraka, Bernieria tenebrosa

This species is poorly known and recorded
from only a few scattered localities on Madagas-
car. During our field survey in the PN de Maro-
jejy, it was observed on one occasion on 16 Oc-
tober, at 0900 hr, in the 775 m zone. A group of
three individuals was seen not far from a stream,
within an area of forest with few large trees and
with areas of open understory vegetation. The
ground was covered by a thick carpet of leaves.
The group was seen foraging on the ground and,
occasionally, climbing up to 0.5 m on low branch-
es, when calling. The call was a single or repeated
"tsit," like other small members of this genus.
Their vocalizations were recorded, and subse-
quent play-backs brought the birds to approxi-
mately 1 m from the observer. This species was
not previously reported for the Marojejy Massif
and is apparently uncommon in the reserve. Its

194

FIELDIANA: ZOOLOGY

northern limit was previously thought to be the
Maroantsetra region (Langrand, 1995).

On the basis of the published literature, the
Dusky Tetraka was known only from a small
number of museum specimens and from sight ob-
servations from the region of Andasibe, the Si-
hanaka Forest, and Maroantsetra (Langrand,
1990). More-recent records of this species are
from lowland forest around the Baie d'Antongil
(Langrand, 1995), Zahamena (Randriamanindry,
1995), and Analamazaotra (Langrand, 1995). This
species was not recorded on any of the previous
World Wide Fund for Nature (WWF) elevational
transects (PN d'Andringitra, RS d'Anjanaharibe-
Sud, or PN d'Andohahela).

Gray-crowned Tetraka, Bernieria cinereiceps

In the Marojejy forest, this species was ob-
served within an elevational band from 1125 to
1675 m. It was not detected frequently on point
counts, perhaps largely because of its rather weak
calls (Hawkins et al., 1998). Twelve individuals
were captured in mist nets, evenly distributed be-
tween these two elevational zones (Table 10-6).
In the 1 250 m zone, this species was seen feeding
in dense understory habitat; at 1625 m, it was ob-
served on several occasions in mixed-species
flocks. At numerous localities across the moun-
tains of eastern Madagascar, this species is com-
mon in montane habitat, generally at elevations
above 1000 m or so.

Madagascar Yellowbrow, Bernieria
xanthophrys

This species was recorded in the 1250 m and
1625 m zones. A small group was found in dense
primary forest vegetation, where they were ob-
served calling along a shaded valley. Safford and
Duckworth (1990) found this species to be com-
mon between 1200 and 1400 m in the same sector
of the reserve.

As with several other species of small passer-
ines found in the montane forest zone of eastern
Madagascar, this species was previously thought
to be fairly rare (e.g., Langrand, 1995). However,
with recent exploration of the montane regions, it
is now known to occur at many localities between
the Anjanaharibe-Sud Massif (Hawkins et al.,
1998) and the Andohahela Massif (Goodman et
al., 1997b).

(Pollen's Vanga, Xenopirostris polleni

This species has been tentatively identified
from the Marojejy region [Benson et al., 1977],
but it was not recorded in the reserve either during
the 1988 Cambridge mission [Safford & Duck-
worth, 1990] or during our 1996 field trip to the
site. Further, this species was not recorded in the
nearby RS d' Anjanaharibe-Sud [Hawkins et al.,
1998]. It appears to be more widespread in the
southern Malagasy eastern forest, particularly in
the vicinity of the PN d'Andohahela [Goodman
et al., 1997b], than in the northern portion of the
island.)

Bernier's Vanga, Oriolia bernieri

During our field observations in the PN de Ma-
rojejy, Bernier's Vanga was recorded only in the
lowland forest between 325 and 600 m. The latter
elevation was on a ridge along the trail between
the 450 and 775 transect zones. O. bernieri ap-
pears to be uncommon to rare in the area of the
Marojejy forest we visited. However, it is fairly
common in the southwestern portion of the re-
serve, at approximately 800 m (Safford & Duck-
worth, 1990). In the RS d' Anjanaharibe-Sud, it
was observed once, at 900 m, during nearly 2
months of field work (Hawkins et al., 1998).

Helmet Vanga, Euryceros prevostii

The majority of records of the Helmet Vanga
are from northeastern Madagascar, including the
region of the Marojejy and Anjanaharibe-Sud
Massifs (Griveaud, 1960; Hawkins et al., 1998).
During the 1996 mission to this site, the Helmet
Vanga was recorded between 400 and 1675 m.
One individual was captured in a mist-net at 445
m (Table 10-6).

Close observations were made of the foraging
habits of a Helmet Vanga. This bird captured in-
sects while in flight and then perched 4-6 m
above the ground to consume the prey. After this
feeding bout, it flew to a nearby stream, wetted
its feathers, and then flew to a branch to dry off.
Not long after that, the bird was observed feeding
on the ground, foraging just alongside an individ-
ual of Bernieria madagascariensis. Euryceros ap-
pears to be widely distributed within the Marojejy
forest. In the RS d' Anjanaharibe-Sud, it was fairly
common at 875 m and was recorded once at 1260
m (Hawkins et al., 1998).

GOODMAN ET AL.: BIRDS

195

Common Myna, Acridotheres tristis

This species, introduced to Madagascar, has not
yet been recorded in the PN de Marojejy (Nicoll
& Langrand, 1989; Safford & Duckworth, 1990;
present study). However, it was noted in 1996,
along the main road not far from Manantenina,
and is common in the Andapa Basin.

Conservation Considerations

In recent years, with the expansion of the num-
ber of conservation projects in the world, great
emphasis has been placed on the importance of
reserves having high diversity and high levels of
endemic species. Numerous systems have been
devised to validate and quantify these variables or
to measure levels of threat to organisms (e.g.,
Mace & Lande, 1991; Mace et al., 1992). In a
world in which the remaining tropical forests are
disappearing before us, such a system of prioriti-
zation is perhaps obligatory. We would like to
point out that these systems are often based on
insufficient data, and the outcome is often an em-
phasis on species rather than on habitat. In turn,
these values often reflect a misrepresentation of
the importance of certain areas or, in another
sense, may diminish the conservation values of
other, lesser known sites.

In 1994, Collar et al. published an important
document that used new criteria established by the
IUCN for assessment of the threat status of bird
species. The categories, based on or modified
from the system of Mace and Stuart (1994), were
proposed for extant birds. The definitions of these
categories are based on a complicated series of
variables and are too detailed and beyond the
scope of currently available population and de-
mographic information for many species. To help
put these definitions into context, we provide
short summary statements of the categories, based
on Collar et al. (1994, p. 15): (1) critical: "facing
an extremely high risk of extinction in the wild in
the near future"; (2) endangered: "facing a very
high risk of extinction in the wild in the near fu-
ture"; (3) vulnerable: "facing a high risk of ex-
tinction in the wild in the medium-term future";
and (4) near-threatened: not falling into any of the
above categories but "close to qualifying for the
[above] threatened categories." (This category
does not directly follow the Mace et al. classifi-
cation.)

Collar et al. (1994) thoroughly reviewed orni-
thological information available at that time and,
for Madagascar, they classified 5 species as criti-
cal, 5 as endangered, 18 as vulnerable, and 16 as
near-threatened, of which 1, 2, 8, and 10 species,
respectively, are known from the PN de Marojejy
(Table 10-9). On the basis of numerous biological
inventories since the publication of this compila-
tion and the ensuing rapid increase in ornitholog-
ical knowledge on Malagasy birds, a re-evaluation
of these categories is needed.

Eutriorchis astur was previously thought to be
extremely rare and on the verge of extinction.
With a series of recent observations of this species
at several different sites on the island (see p. 192),
it is now known to have a much broader distri-
bution than previously thought. On the basis of
new information, this species should probably be
moved from the critical to the endangered cate-
gory.

The case of Neodrepanis hypoxantha is an ex-
cellent example of how further research has great-
ly elucidated the status of this species (Hawkins
et al., 1997). This bird, which once was known
only from slightly more than a dozen museum
specimens and a handful of observations, has re-
cently been documented as occurring in montane
and sclerophyllous forest from the Marojejy and
Anjanaharibe-Sud Massifs in the north to the An-
dohahela Massif in the far south. It is a species
with a limited altitudinal distribution but a broad
geographic range. At some sites it is common;
density estimates based on point counts yielded
up to several thousand individuals per square ki-
lometer, although in some cases these may be
overestimates (Hawkins et al., 1998, p. 1 12). This
species should probably be removed from the
near-threatened list.

Several other species previously classified as
either vulnerable (Mesitornis unicolor, Brachyp-
teracias leptosomus, B. squamiger, Atelornis
crossleyi, Bernieria cinereiceps, B. xanthophrys,
Xenopirostris polleni, and Oriolia bernieri) or
near-threatened (Accipiter madagascariensis, A.
henstii, Atelornis pittoides, Euryceros prevostii,
Monticola sharpei, Dromaeocercus brunneus,
Randia pseudozosterops, and Pseudobias wardi)
are all more common and broadly distributed than
previously thought. Further, several of these spe-
cies are known to live and nest in degraded hab-
itats (Table 10-9) and occur in relatively high den-
sities (Table 10-8). Examples from forests outside
the PN de Marojejy include Atelornis crossleyi,
with more than 65 individuals per square kilo-

196

FIELDIANA: ZOOLOGY

Table 10-9. Forest birds recorded in the PN de Marojejy that were considered threatened by Collar et al. (1994),
and their current status.

Threat

Current

Species

Category

codes'

status 2

Eutriorchis astur

Critical

1,2,3,5

A, B

Neodrepanis hypoxantha

Endangered

B,C

Bernieria tenebrosa

Endangered

D

Mesitornis unicolor

Vulnerable

B

Brachypteracias leptosomus

Vulnerable

B,E

Brachypteracias squamiger

Vulnerable

1,2,4

B

Atelornis crossleyi

Vulnerable

B,C, E

Bernieria cinereiceps

Vulnerable

B,C, E

Bernieria xanthophrys

Vulnerable

B,C, E

Newtonia fanovanae

Vulnerable

A, D

Xenopirostris polleni

Vulnerable

B,E

Oriolia bernieri

Near-threatened

B

Lophotibis cristata

Near-threatened

F

Accipiter madagascariensis

Near-threatened

B

Accipiter henstii

Near-threatened

B,F

Atelornis pittoides

Near-threatened

B, E

Euryceros prevostii

Near-threatened

B

Monticola sharpei

Near-threatened

B, C, E, F

Hartertula flavoviridis

Near-threatened

B

Dromaeocercus brunneus

Near-threatened

B,C, E

Randia pseudozosterops

Near-threatened

B, E

Pseudobias wardi

Near-threatened

B, E

1 Key to threat codes (Collar et al., 1994): 1 = loss or alteration of habitat; 2 = hunting, persecution (including
accidental trapping), egg collecting (subsistence); 3 = disturbance (by humans, stock); 4 = introduced species (pred-
ators, competitors, herbivores, diseases); 5 = small range or population. Threat codes were not provided for near-
threatened species.

2 Key to current status: A = rediscovered after a long hiatus of information on this species in the wild; B = now
known to have a much broader range than previously surmised; C = occurs in altitudinally limited but geographically
widespread habitat; D = still poorly known, but apparently more widespread than previously thought; E = in some
areas a common forest inhabitant; F = occurs in disturbed habitat with moderate to considerable human pressure.

meter, Monticola sharpei, with more than 140 in-
dividuals per square kilometer, and Dromaeocer-
cus brunneus, with almost 50 individuals per
square kilometer (Hawkins et al., 1998; Hawkins
& Goodman, 1999). One enigmatic species, Ber-
nieria tenebrosa, remains poorly known, and its
"threatened" status should remain unchanged.

Recent biogeographic analyses of the distribu-
tion of forest-dependent species in the eastern hu-
mid forest has shown that most species are dis-
tributed widely and that the avifauna across this
vast region is relatively homogeneous (Wilme,
1996; Goodman et al., 1997b), although there is
a decrease in bird species richness from north to
south (see p. 188 for further discussion of this
point). The high conservation status allocated to
the PN de Marojejy by Collar and Stuart (1988)
and Evans et al. (1992) is a direct result of the
site's rich avifauna; however, this is also one of
the more intensively studied mountains in the or-
nithologically rich northern portion of the island.

Almost certainly, other sites in the north that in-
clude a broader elevational range of intact forests,
from the lowlands to above tree line, hold rich
avifaunas equal to that known from the PN de
Marojejy. Several such sites, such as the RNI de
Tsaratanana and adjoining forest blocks, include
vast expanses of relatively intact forest and are of
key biogeographic importance, since the vegeta-
tional structure on the east and west slopes are
distinctly different. Thus, in light of these points
and the fact that Marojejy is already part of an
extensive conservation program, future conser-
vation work should emphasize nearby forested ar-
eas, particularly those that are connected to one
another. An excellent example is the forest of Be-
taolana, a forested corridor that connects the PN
de Marojejy and the RS d'Anjanaharibe-Sud, and
other forested corridors in the north that link var-
ious massifs and large forest blocks. These for-
ested corridors are presumably critical for move-
ments and genetic exchange between populations,

GOODMAN ET AL.: BIRDS

197

and the corridors should be given high priority for
research and conservation status. The remaining
lowland forest within and outside these reserves
is the most threatened habitat type in the region,
and conservation programs should pay particular
attention to amelioration of the human pressures
on this biotype.

Several other regions of the eastern humid forest
that are not part of the current protected areas of
the island have been identified as biologically poor-
ly known and of probable high biotic importance
(Ganzhom et al., 1997). Further south, these sites
include large areas of lowland forest. Examples of
these forests include Rantabe, the "Sihanaka For-
est" southeast of Lac Alaotra, and between Didy
and Fito, the corridor between the PN de Rano-
mafana and the PN d'Andringitra, including the
forests of Tolongoina and Ikongo. The Sihanaka
Forest is an interesting case in point. Collar and
Stuart (1988) gave this site the highest conserva-
tion status of any forest on the island, although this
zone is largely unknown in modern times, and all
current information is based largely on extensive
collections made at the site more than 50 years ago.
Until the recent wave of ornithological activity on
Madagascar, such old collections represented a
considerable proportion of the available data on the
birds of the island. We now know that, although
such information is extremely important, it does
not represent the true distribution of the vast ma-
jority of forest-dwelling species.

Finally, the recent increase in inventory work in
the eastern humid forest of Madagascar has been
conducted largely in areas that are already protect-
ed. Given that baseline data exist for many such
sites and that human pressure is higher outside
rather than inside protected areas, now is the crit-
ical time to document the biota existing outside the
reserves, to clearly set the priorities for future con-
servation on the island. In the northern portion of
the island, certain reserves and surrounding forests
are poorly known. An excellent example is the RNI
de Tsaratanana, which contains the highest moun-
tain on the island and extensive areas of forest.
Little is known about the biota of this site, and only
the drier western slope of the mountain is within
the reserve. The regions on the eastern portion of
the massif are almost certainly of critical biological
importance but are in need of baseline data from
biological inventories as well as assessments of hu-
man pressures.

These arguments are by no means meant to de-
tract from the importance of the PN de Marojejy
for safeguarding the high and remarkable biodi-

versity that the reserve holds. As clearly men-
tioned by Evans et al. (1992), the extensive low-
land areas of the reserve have experienced severe
deforestation, and these pressures continue. Many
key species (sensu Evans et al. [1992]) are re-
stricted to these habitats. On the other hand, an-
other suite of their key bird species occurs in the
upper montane and sclerophyllous forests, and
they are under little threat associated with habitat
loss. These forest types are still relatively exten-
sive although not well represented in the protected
areas system (Du Puy & Moat, 1996). The factors
that have led to high human pressure on montane
forest at sites such as the PN d'Andringitra par-
ticularly easy access, local cultural traditions, and
vast zones for cattle pasturage (Berner, 1997)
do not exist in the PN de Marojejy. Thus, these
forest types are not under any serious threat in the
PN de Marojejy.

One of the proposed reasons to change the sta-
tus of the Marojejy Massif from Reserve Natu-
relle Integrate (with highly restricted access) to
Pare National was to allow tourist access to the
site (see Chapter 1). Thus, in principle, an influx
of revenue divided among people living around
the reserve would result in improvement of their
financial means and, at the same time, reduce their
need to exploit forest products or to use slash-and-
burn agricultural practices for subsistence. In the
final assessment, the PN de Marojejy contains an
important number of endemic bird species.
Among the currently protected areas of Madagas-
car, it has tremendous possibilities for attracting
ecotourism and continued scientific investigations,
and the avifauna of the massif will be one of the
main features to sustain this interest.

Acknowledgments

We are grateful to the Association Nationale
pour la Gestion des Aires Protegees and the Min-
istere des Eaux et Forets for permission to conduct
this study. The staff of WWF Andapa helped us
considerably with logistical and technical prob-
lems. We continue to be indebted to M. Mamy Ra-
vokatra for his logistic and scientific aid (including
the mist-netting operations) during our field mis-
sions. Tom Schulenberg provided extensive com-
ments on an earlier version of this chapter.

198

FIELDIANA: ZOOLOGY

Literature Cited

Association Nationale pour la Gestion des Aires
Protegees. 1998. Synthese de Recherches dans le Re-
seau d' Aires Protegees de Madagascar. Association
Nationale pour la Gestion des Aires Protegees, Anta-
nanarivo, 219 pp.

Benson, C. W., J. F. R. Colebrook-Robjent, and A.
Williams. 1976. Contribution a l'ornithologie de
Madagascar. L'Oiseau et Revue Francaise
d'Ornithologie, 46: 103-134, 209-242, 367-386.

. 1977. Contribution a l'ornithologie de Mada-
gascar. L'Oiseau et Revue Francaise d'Ornithologie,
47: 41-61, 167-191.

Berner, P. O. 1997. Les forets de transition de la region
Nord-Est du massif de l'Andringitra, Madagascar:
Monitoring d' impacts anthropiques et proposition
d'amenagement. Akon'ny Ala, 21: 9-26.

Bibby, C. J., N. D. Burgess, and D. A. Hill. 1992. Bird
Census Techniques. Academic Press, London, 257 pp.

Collar, N. J., M. J. Crosby, and A. J. Stattersfield.
1994. Birds to Watch, 2: The World List of Threatened
Birds. Bird Life International, Cambridge, 407 pp.

Collar, N. J., and S. N. Stuart. 1985. Threatened
Birds of Africa and Related Islands: The ICBP/IUCN
Red Data Book, part I, 3rd ed. International Council
for Bird Preservation and International Union for the
Conservation of Nature and Natural Resources, Cam-
bridge, United Kingdom.

. 1988. Key Forests for Threatened Birds in Af-
rica. International Council for Bird Preservation,
Monograph no. 3, Cambridge, United Kingdom.

Du Puy, D. J., and J. Moat. 1996. A refined classifi-
cation of the primary vegetation of Madagascar based
on the underlying geology: Using GIS to map its dis-
tribution and to assess its conservation status, pp.
205-218. In Lourenco, W. R., ed., Biogeographie de
Madagascar. ORSTOM Editions, Paris, 588 pp.

Evans, M. I. 1991. The Red-tailed Newtonia Newtonia
fanovanae in the Ambatovaky Reserve, north-east
Madagascar. Bird Conservation International, 1: 47-
52.

Evans, M. I., J. W Duckworth, A. F. A. Hawkins, R.
J. Safford, B. C. Sheldon, and R. J. Wilkinson.
1992. Key bird species of Marojejy Strict Nature Re-
serve, Madagascar. Bird Conservation International,
2: 201-220.

Fjeldsa, J., S. M. Goodman, T S. Schulenberg, and
B. Slikas. 1999. Molecular evidence for relationships
of Malagasy birds, pp. 3084-3094. In Adams, N., and
R. Slotow, eds., Proceedings of the 22nd International
Ornithological Congress. University of Natal, Durban.

Ganzhorn, J. U., B. Rakotosamimanana, L. Hannah,
J. Hough, L. Iyer, S. Olivieri, S. Rajaobelina, C.
Rodstrom, and G. Tilkin. 1997. Priorities for biodi-
versity conservation in Madagascar. Primate Report,
Special Issue, 48-1: 1-81.

Goodman, S. M., A. Andrianarimisa, L. E. Olson, and
V. Soarimalala. 1996. Patterns of elevational distri-
bution of birds and small mammals in the humid for-
ests of Montagne d'Ambre, Madagascar. Ecotropica,
2: 87-98.

Goodman, S. M., and J. U. Ganzhorn. 1997. Rarity of

figs (Ficus) on Madagascar and its relationship to a
depauperate frugivore community. Revue d'Ecologie,
52: 321-329.

Goodman, S. M., A. E A. Hawkins, and C. A. Do-
mergue. 1997a. A new species of vanga (Vangidae,
Calicalicus) from southwestern Madagascar. Bulletin
of the British Ornithologists' Club, 117: 4-10.

Goodman, S. M., O. Langrand, and B. M. Whitney.
1995. A new genus and species of passerine from the
eastern rainforest of Madagascar. Ibis, 138: 153-159.

Goodman, S. M., and P. Parrillo. 1997. A study of the
diets of Malagasy birds based on stomach contents.
Ostrich, 68: 104-113.

Goodman, S. M., M. Pidgeon, A. F A. Hawkins, and
T S. Schulenberg. 1997b. The birds of southeastern
Madagascar. Fieldiana: Zoology, n. s., 87: 1-132.

Goodman, S. M., and M. S. Putnam. 1996. The birds
of the eastern slopes of the Reserve Naturelle Integrale
d'Andringitra, Madagascar, pp. 171-190. In Goodman.
S. M., ed., A Floral and Faunal Inventory of the Eastern
Slopes of the Reserve Naturelle Integrale d'Andringitra,
Madagascar: With Reference to Elevational Variation.
Fieldiana: Zoology, n.s., 85: 1-319.

Goodman, S. M., and B. Rasolonandrasana. In press.
Elevational zonation of birds, insectivores, rodents,
and primates on the slopes of the Andringitra Massif,
Madagascar. Journal of Natural History.

Goodman, S. M., and T S. Schulenberg. 1991. The
rediscovery of the Red-tailed Newtonia Newtonia fa-
novanae in south-eastern Madagascar with notes on
the natural history of the genus Newtonia. Bird Con-
servation International, 1: 33-45.

Goodman, S. M., and L. E. Weight. In press. The ge-
neric and species relationships of the reputed endemic
Malagasy genus Pseudocossyphus (family Turdidae).
Ostrich.

Griveaud, P. 1960. Une mission entomologique au Ma-
rojejy. Naturaliste Malgache, 12: 43-55.

Hawkins, A. F A. 1999. Altitudinal and latitudinal dis-
tribution of east Malagasy forest bird communities.
Journal of Biogeography. 26: 447-458.

Hawkins, A. E A., and S. M. Goodman. 1999. Bird
community variation with elevation and habitat in par-
cels 1 and 2 of the Reserve Naturelle Integrale
d'Andohahela, Madagascar, pp. 175-186. In Good-
man, S. M., ed., A Floral and Faunal Inventory of
Reserve Naturelle Integrale d'Andohahela, Madagas-
car: With Reference to Elevational Variation. Fieldi-
ana: Zoology, n.s., 94: 1-297.

Hawkins, A. F. A., R. J. Safford, J. W Duckworth,
and M. I. Evans. 1997. Field identification and status
of the sunbird asities Neodrepanis of Madagascar.
Bulletin of the African Bird Club, 4: 36-41.

Hawkins, A. E A, J.-M. Thiollay, and S. M. Goodman.
1998. The birds of the Reserve Speciale
d'Anjanaharibe-Sud, Madagascar, pp. 43-147. In
Goodman, S. M., ed., A Floral and Faunal Inventory
of the Reserve Speciale d'Anjanaharibe-Sud, Mada-
gascar: With Reference to Elevational Variation. Fiel-
diana: Zoology, n.s., 90: 1-246.

Hawkins, A. F A., A. Andrianarimisa, O. M. Rako-
tonomenjanahary, and V Raminoarisoa. In press.

GOODMAN ET AL.: BIRDS

199

Inventaire des oiseaux dans la Reserve Naturelle In-
tegrate de Zahamena. Ostrich.

Jenkins, M. D.. ed. 1990. Madagascar: Profil de
rEnvironnement. UICN/PNUEAVWF. Gland et Cam-
bridge. 439 pp.

Karr. J. R. 1981. Surveying birds with mist nets. Stud-
ies in Avian Biology, 6: 62-67.

KOECHLIN, J., J.-L. GUILLAUMET. AND P. MORAT. 1974.

Flore et Vegetation de Madagascar. J. Cramer, Vaduz,
629 pp.

Inskipp. C. and T. Inskipp. 1991. Birds of Nepal, 2nd
ed. Christopher Helm, London.

Langrand, O. 1990. Guide to the Birds of Madagascar.
Yale University Press. New Haven. 364 pp.

. 1995. Guide des oiseaux de Madagascar. De-

lachaux et Niestle. Lausanne. 415 pp.

Mace. G., N. Collar. J. Cooke. K. Gaston. J. Gins-
burg. N. Leader-Williams. M. Maunder, and E. J.
Milner-Gulland. 1992. The development of new cri-
teria for listing species on the IUCN Red List. Spe-
cies, 19: 16-22.

Mace, G. M., and R. Lande. 1991. Assessing extinction
threats: Toward a reevaluation of IUCN threatened
species categories. Conservation Biology. 5: 148-157.

Mace, G. M.. and S. Stuart. 1994. Draft IUCN Red
List categories. Species, 21-22: 13-24.

Maclean. G. L. 1990. Ornithology for Africa. Univer-
sity of Natal Press. Pietermaritzburg.

Morris. P. A. J., and A. F. A. Hawkins. 1998. The Birds
of Madagascar: A Photographic Guide. Pica Press.
East Sussex. United Kingdom. 316 pp.

Nicoll. M. E.. and O. Langrand. 1989. Madagascar:
Revue de la Conservation et des Aires Protegees.
World Wide Fund for Nature. Gland. 374 pp.

Prum, R. O., and V R. Razafindratsita. 1997. Lek
behavior and natural history of the Velvet Asity (Phi-
lepitta castanea: Eurylaimidae). Wilson Bulletin. 109:
371-392.

Rand. A. L. 1932. Mission Franco-Anglo-Americaine a
Madagascar: Notes de Voyage. LOiseau et la Revue
Francaise d'Ornithologie (nouvelle serie). 2: 227-282.

. 1936. The distribution and habits of Madagas-
car birds: A summary of the field notes of the Mission
Zoologique Franco-Anglo-Americaine a Madagascar.
Bulletin American Museum of Natural History. 72:
143-499.

Randriamantndry. J. -J. 1995. The Dusky Greenbul
Phyllastrephus tenebrosus in Zahamena Strict Nature
Reserve. Madagascar. Newsletter of the Working
Group on Birds in the Madagascar Region. 5(2): 5.

Raxworthy. C. J., and P. R. Colston. 1992. Conclusive
evidence for the continuing existence of the Mada-
gascar Serpent-eagle Eutriorchis astur. Bulletin of the
British Ornithologists" Club. 112: 108-111.

Razafindratsita. V R. 1995. Etude biologique et ecol-
ogique de Philepitta castanea (Mullen 1776) son role
dans la regeneration du sois-bois forestier du Pare Na-
tional de Ranomafana. Memoire de Diplome d' Etudes

Approfondies. Universite d' Antananarivo. Antanana-
rivo.

Remsen. J. V. Jr. and D. A. Good. 1996. Misuse of
data from mist-net captures used to assess relative
abundance in bird populations. Auk. 113: 381-398.

Reynolds. R. T. J. M. Scott, and R. A. Nussbaum.
1980. A variable circular-plot method for estimating
bird numbers. Condor. 82: 309-313.

Safford. R. J., and J. W Duckworth, eds. 1990. A
wildlife survey of Marojejy Reserve. Madagascar.
ICBP Study Report 40. International Council for Bird
Preservation. Cambridge.

Schulenberg. T. S., S. M. Goodman, and J.-C. Raza-
fimahaimodison. 1993. Genetic variation in two sub-
species of Nesillas typica (Sylviinae) in south-east
Madagascar, pp. 173-177. In Wilson. R. T, ed. Birds
and the African Environment: Proceedings of the 8th
Pan-African Ornithological Congress. Annales Musee
Roy ale de l'Afrique Centrale (Zoologie). 268.

Sheldon. B. C. and J. W Duckworth. 1990. Redis-
covery of the Madagascar Serpent-eagle Eutriorchis
astur. Bulletin of the British Ornithologists' Club.
110: 126-130.

Sinclair. I., and O. Langrand. 1998. Birds of the In-
dian Ocean Islands. Struik Publishers. Ltd.. Cape
Town, 184 pp.

Stotz. D. E. J. W. Fitzpatrick, T. A. Parker. HI. and
D. K. Moskovits. 1996. Neotropical Birds: Ecology
and Conservation. The University of Chicago Press.
Chicago. 478 pp.

Straka. H. 1996. Histoire de la vegetation de Mada-
gascar oriental dans les derniers 100 millenaires. pp.
37-47. In Lourenco. W R.. ed.. Biogeographie de
Madagascar. Editions ORSTOM. Paris.

Thiollay. J.-M. 1998. The diurnal raptors (Falconifor-
mes) of the Reserve Speciale d'Anjanaharibe-Sud:
Abundance, distribution, and conservation, pp. 129-
138. In Goodman, S. M.. ed., A Floral and Faunal
Inventory of the Reserve Speciale d'Anjanaharibe-
Sud. Madagascar: With Reference to Elevational Var-
iation. Fieldiana: Zoology, n.s.. 90: 1-246.

Thompson. P. M.. and M. I. Evans. 1992. The threat-
ened birds of Ambatovaky Special Reserve. Mada-
gascar. Bird Conservation International, 2: 221-237.

Thorstrom. R.. and R. T. Watson. 1997. Avian inven-
tory and key species of the Masoala Peninsula. Mad-
agascar. Bird Conservation International. 7: 99-115.

Thorstrom. R.. R. Watson. B. Damary, E Toto. M.
Bab. and V Baba. 1995. Repeated sightings and re-
cent capture of a live Madagascar Serpent-eagle Eu-
triorchis astur. Bulletin of the British Ornithologists"
Club. 115: 40-45.

Wilme. L. 1996. Composition and characteristics of bird
communities in Madagascar, pp. 349-362. In Lour-
enco. W. R.. ed.. Biogeographie de Madagascar. OR-
STOM Editions. Paris. 588 pp.

Yamagishi. S.. T. Masuda. and H. Rakotomanana.
1997. [A Field Guide to the Birds of Madagascar.] [In
Japanese and Malagasy.] Kaiyusha Publishers. Ltd..
Tokyo. 160 pp.

200

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Chapter 11

Tenrecs (Lipotyphla: Tenrecidae) of the Pare National
de Marojejy, Madagascar

Steven M. Goodman 1 and Paulina D. Jenkins 2

Abstract

Tenrecs (Lipotyphla: Tenrecidae) were studied in northeastern Madagascar in the Pare Na-
tional (PN) de Marojejy in five zones across a forested elevational gradient from 450 to 1875
m. The transect went through a variety of habitats, including lowland (450-775 m), montane
(1225-1625 m), and sclerophyllous/mossy (1875 m) forests. Trapping techniques involved pit-
fall buckets and standard live small mammal traps. Fifteen species of lipotyphlans were re-
corded in the reserve, including 12 species of shrew tenrecs (Microgale). Among Microgale,
there is no clear evidence of species replacement along the elevational gradient surveyed within
the PN de Marojejy. New information is presented on morphology, reproduction, development,
and ecology of the Tenrecidae captured in this reserve.

No species of Tenrecidae was recorded across the complete elevational gradient. The two
most elevationally widespread species were Microgale talazaci and M. parvula, which were
trapped between 450 and 1625 m. Five species occurred within the montane forest zone: M.
dobsoni, M. fotsifotsy, M. gymnorhyncha, M. longicaudata, and M. soricoides, and three species
occurred across the montane and sclerophyllous/mossy forest habitats: M. cowani, M. monti-
cola, and M. principula. Only two species were captured in traps at a single elevational zone:
M. brevicaudata at 475 m and M. gracilis at 1875 m. The only species with an apparently
disjunct altitudinal distribution was the mole tenrec Oryzorictes hova, which was collected in
lowland forest at 450 m and in montane forest at 1625 m; this result is almost certainly an
artifact of the difficulty in trapping members of this genus.

The greatest Tenrecidae species richness was at 1625 m, where nine species (including eight
species of Microgale) were found. The 775 m and 1875 m zones had the lowest species
richness, with only four species each. The highest densities of lipotyphlans in the reserve, as
measured by pitfall trap success, were in the montane forest, which across the different pitfall
lines averaged 19.5% in the 1225 m zone (range 14.3%-24.7%) and 15.6% in the 1625 m
zone (range 11.7%-19.5%). The lowland forests at 450 and 775 m had the lowest Tenrecidae
capture rates; 3.8% at 450 m and 1.9% at 775 m.

A comparison is made between the Tenrecidae found in the PN de Marojejy and those
captured by means of identical trapping techniques along an elevational transect in the Reserve
Speciale (RS) d' Anjanaharibe-Sud, 40 km to the west. The vast majority of species were found
in common between these two reserves (Oryzorictes hova, Microgale cowani, M. dobsoni, M.
fotsifotsy, M. gymnorhyncha, M. longicaudata, M. monticola, M. parvula, M. principula, M.
soricoides and M. talazaci), but there are a few differences, with M. brevicaudata and M.
gracilis obtained only in PN de Marojejy and M. dryas only in the RS d' Anjanaharibe-Sud.
The altitudinal distributions of these species in the two reserves are broadly similar.

1 Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, U.S.A.

2 The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom.

GOODMAN & JENKINS: LIPOTYPHLA 201

Resume

Les Tenrecs (Lipotyphla: Tenrecidae) ont ete etudies au nord-est de Madagascar dans le Pare
National (PN) de Marojejy ou les cinq zones d'etudes retenues traversaient un gradient alti-
tudinal de 450 a 1875 m. Le transect parcourait une variete d'habitats incluant la foret de basse
altitude (450-775 m), de montagne (1225-1625 m), et sclerophylle/de mousses (1875 m). Les
techniques de piegeage comprenaient les lignes de seaux enterres et les pieges standard pour
la capture de petits mammiferes vivants. Quinze especes des lipotyphlans ont ete relevees dans
la reserve, y compris 12 especes de tenrec-musaraignes (Microgale) pour lesquels aucune ev-
idence claire n'a permis de montrer un remplacement d'especes le long du gradient altitudinal
examine dans le PN de Marojejy. Les nouvelles informations relatives a la morphologie, la
reproduction, le developpement, et l'ecologie des Tenrecidae captures dans cette reserve sont
ici presentees.

Aucune espece de Tenrecidae n'a ete enregistree sur l'ensemble du gradient altitudinal. Les
deux especes les plus largement distributes sur le gradient etaient Microgale talazaci et M.
pan'ula, capturees a tous les niveaux entre 450 et 1625 m d'altitude. Cinq especes ont ete
rencontrees dans la zone de foret de montagne: M. dobsoni, M. fotsifotsy, M. gymnorhyncha,
M. longicaudata et M. soricoides, et trois especes a travers les habitats de la foret de montagne
et la foret sclerophylle/de mousse: M. cowani, M. monticola, et M. principula. Seules deux
especes etaient limitees a une seule zone altitudinale: M. brevicaudata a 475 m and M. gracilis
a 1875 m.

La richesse specifique la plus importante a ete relevee a 1625 m d'altitude avec 9 especes
(y compris 8 especes de Microgale). Dans les zones a 775 m et 1875 m, aura ete relevee la
plus basse richesse specifique avec quatre especes. Les densites de lipotyphlans les plus elevees
de la reserve, mesurees par le succes de la ligne de piegeage employant les baquets, ont ete
mesurees dans la foret de montagne avec une moyenne, calculee sur les differentes lignes, de
19,5% dans la zone a 1225 m (ecart 14,3-24,7%) et 15,6% dans la zone a 1625 m (ecart 11,
7-19,5%). Le plus faible taux de capture de Tenrecidae a ete releve dans foret de basse altitude
a 450 et 775 m: 3,8% a 450 m et 1,9% a 775 m.

Une comparaison est faite entre les Tenrecidae releves dans le PN de Marojejy et ceux
captures en utilisant des techniques de piegeage identiques le long d'un transect altitudinal
d'elevation dans le Reserve Speciale (RS) d'Anjanaharibe-Sud, situee 40 kilometres a l'ouest.
La grande majorite des especes sont communes aux deux reserves (Oryzorictes hova, Microgale
cowani, M. dobsoni, M. fotsifotsy, M. gymnorhyncha, M. longicaudata, M. monticola, M. par-
vula, M. principula, M. soricoides, et M. talazaci), mais il y a quelques differences dont M.
brevicaudata et M. gracilis qui n'ont ete releves que dans le PN de Marojejy et M. dryas que
dans la RS d'Anjanaharibe-Sud. La distribution altitudinale de ces especes est globalement
semblable dans les deux reserves.

Introduction

In 1987 MacPhee published an important re-
vision of Microgale, the most speciose genus of
Tenrecidae, that provided the critical foundation
for more recent studies on the species limits and
biogeography of this group. One of the problems
that plagued advancement of tenrecid systematics
was the lack of comparative material, particularly
specimens with intact skulls and indicators of ex-
ternal measurements, sex, and provenance.
MacPhee (1987) amassed the Microgale material
held in virtually all the major museums of the

world but was able to study only approximately
120 specimens. Faced with sorting out synony-
mies and species limits for 22 named taxa, he had
an average of only 5.5 specimens per taxon.
Clearly, such sample sizes are insufficient for un-
derstanding aspects of age, sexual, and geographic
variation within a species.

Over the course of the past decade there has
been a resurgence in field efforts on Madagascar,
and a portion of this work has centered on bio-
logical inventories in poorly known areas of for-
est. A considerable amount of new small mammal
material has been obtained. These new collec-
tions, combined with further research on existing

202

FIELDIANA: ZOOLOGY

specimens in museums, have resulted in the de-
scription of numerous new species of Malagasy
insectivores and rodents and have allowed mam-
malogists to completely revamp ideas about the
diversity, species limits, and biogeography of
these animals on Madagascar. For Microgale
alone, the collections now available to help clarify
the relationships of this group are many times
greater than what was available for MacPhee's re-
vision. Further, this new material, combined with
preserved tissues for biochemical analyses, has
provided the means to commence studies on as-
pects of the phylogeny and relationships of the
Tenrecidae.

As part of these surveys of poorly known areas
of Malagasy forest, a group of biologists con-
ducted an inventory in late 1996 of the Pare Na-
tional (PN) de Marojejy in the northeastern por-
tion of the island (see Chapter 1). This survey,
organized by World Wide Fund for Nature in the
context of a conservation project in this reserve,
was the fourth annual survey to examine the dis-
tribution of biota along elevational gradients in
eastern humid forests, and, during all of these sur-
veys, concerted efforts have been made to study
Tenrecidae.

Thirteen species of oryzorictine tenrecs were
collected during the 1996 survey of the PN de
Marojejy: Microgale brevicaudata, M. cowani, M.
dobsoni, M. fotsifotsy, M. gracilis, M. gymnorhyn-
cha, M. longicaudata, M. monticola, M. parvula,
M. principula, M. soricoides, M. talazaci, and
Oryzorictes hova. In this chapter we present de-
tails of synonymy, measurements, key characters
for identification, reproduction and population
structure, and distribution for the various species
captured in the reserve. We follow the basic for-
mat used in previous, similar surveys of Malagasy
lipotyphlans (Jenkins et al., 1996; Goodman et al.,
1996a, 1999a; Goodman & Jenkins, 1998). Fur-
thermore, we provide a more comprehensive de-
scription for Microgale brevicaudata, a poorly
known species for which only five specimens, in-
cluding the holotype, were available for Mac-
Phee's (1987) study.

One of the mountains surveyed during the past
few years in a manner comparable to the work
conducted in the PN de Marojejy is the Reserve
Speciale (RS) d'Anjanaharibe-Sud. These two
mountains are 40 km from one another, on op-
posite sides of the Andapa Basin (Fig. 11-1).
Throughout this chapter we compare the results
of tenrecid distribution, density, and species rich-
ness between these two sites, which are at nearly

the same latitude. Information is now available on
the shrew tenrecs occurring at five different
mountains in the eastern part of the country, span-
ning the complete latitudinal gradient of the island
(12E-25E S; Fig. 11-1). In this chapter we also
examine trends in species richness, elevational zo-
nation, and geographic distribution related to this
gradient. Finally, we analyze variation in species
distribution and provide details of altitudinal and
ecological separation.

Materials and Methods

Traplines

The principal technique used to capture lipo-
typhlans consisted of pitfall traps with drift fenc-
es. In each of the elevational zones surveyed (450,
775, 1225, 1625, and 1875 m) separate pitfall
lines were installed in three different topographic
settings (valley bottom, slope, and ridge crest) to
assess possible variation in microhabitat use by
lipotyphlans. Each line was 100 m long and con-
sisted of 1 1 buckets (275 mm deep, 290 mm top
internal diameter, 220 mm bottom internal diam-
eter), 10 m apart, in operation for a minimum of
7 nights. (The only exception was the lines at
1875 m, which were in place for 5 nights.) Small
holes (2 mm diameter) were drilled in the bottom
of each bucket to allow water drainage. Buckets
were sunk to a depth at which the rim was even
with ground level. A barrier (drift fence) made
from plastic sheeting (0.5 m high and 100 m long)
was stapled in a vertical position to thin wooden
stakes. The drift fence bisected all of the buckets
in the line (Voss & Emmons, 1996 [Fig. 7]). A
flange of approximately 50 mm at the bottom of
the standing plastic fence was covered with soil
and leaf litter to prevent animals from moving un-
der the barrier. Lipotyphlans were also captured
with standard Sherman live traps. The trap types,
placement, baits, and other details for these lines
are described in Chapter 12.

Traps and pitfalls were visited at least twice per
day, once at dawn and again in the late afternoon,
and captured animals were removed. A "trap-
day" and a "bucket-day" are defined as a 24-hour
period of use of one of these devices (dawn to
dawn). After heavy rain, water was removed from
the traps by means of a sponge. The survey of the
PN de Marojejy was conducted between 5 Octo-
ber and 19 November 1996, a season during

GOODMAN & JENKINS: LIPOTYPHLA

203

Elevation
- 500 m

500 -1000 m

1000 -1500 m

1500 -2000 m

> 2000 m

Fig. 11-1. Map of mountains for which detailed information is available for the distribution of Lipotyphla along
elevational gradients, and various other sites mentioned in the text.

which all Tenrecidae, including those that are
known to hibernate or aestivate, are expected to
be active (Stephenson, 1994). However, the re-
sults of previous surveys conducted during this
same calendar period seem to indicate that some
of the large spiny tenrecs (Tenrec ecaudatus,
Hemicentetes spp., and Setifer setosus) may not
be very active, or at least that population cycles
are at a low before reproduction, associated with
the upcoming rainy season (Goodman et al.,
1996a; Goodman & Jenkins, 1998).

The majority of captured animals were pre-
pared as standard museum skins with associated

skulls and skeletons, as fluid-preserved carcasses,
or as full skeletons. Voucher specimens will be
divided equally between the Field Museum of
Natural History (FMNH), Chicago, and the De-
partement de Biologie Animale, Universite
d' Antananarivo (UA). Specimens deposited im-
mediately after the survey in the latter institution
have not yet been catalogued and are referenced
individually by the collector's field numbers (UA-
SMG). Animals not kept as voucher specimens
were first marked by clipping a small area of fur
on their backs and then released near the original
trap site. This marking technique allowed first-

204

FIELDIANA: ZOOLOGY

time and previously captured animals to be dif-
ferentiated.

Measurements

Crania were measured by means of digital cal-
ipers and a microscope measuring stage. Dental
nomenclature follows Mills (1966), Swindler
(1976), Butler and Greenwood (1979), MacPhee
(1987), and Jenkins et al. (1996, 1997). Dental
notations are given in parentheses in the text; pre-
maxillary and maxillary teeth are denoted by up-
percase and mandibular teeth by lowercase letters,
as follows: incisor (I/i), canine (C/c), premolar (P/
p), and molar (M/m). A prefix "d" indicates de-
ciduous teeth; thus, dI3 refers to the deciduous
third upper incisor. The following measurements
were made either from specimens in the flesh or
from prepared crania. Abbreviations and defini-
tions for these measurements (all in millimeters,
with the exception of weight, in grams) are as
follows:

BB (breadth of braincase): the greatest distance
measured across the squamosals.

CIL (condyloincisive length): cranial length
from the anterior surface of the first upper
incisor to the articular surface of the occip-
ital condyle.

E (ear length): measured from the notch at the
base of the ear to the distal-most edge of the
pinna.

HB (head and body length): measured from the
tip of the nose to the distal-most point of
the body (at base of tail).

HF (hind foot length): measured from the back
edge of the heel to the tip of the longest toe
(not including claw).

TL (tail length): measured from the base of the
tail (at right angles to the body) to the end
of the distal-most vertebra. Does not include
terminal hair tufts.

UTL (upper tooth-row length): measured from the
anterior surface of the first upper incisor to
the posterior surface of the third upper mo-
lar, parallel to the long axis of the skull.

WT (weight): measured in grams by means of
Pesola spring scales. Animals weighing less
than 10 g were weighed within 0.2 g; those
weighing 10-100 g were weighed within
0.5 g.

Reproductive condition was recorded for males
as length X width of the testes and degree of con-

volution of the epididymides. Females were noted
as nonperforate or perforate and as nonparous or
parous, and the number and location of any em-
bryos and placental scars were recorded. The
mammary formula is presented as the number of
paired axial, abdominal, and inguinal nipples.
The following age classes are recognized:

"Infant": Individuals in which the deciduous an-
temolar dentition and the molars are not fully
erupted and premaxillary, parietal, and basioc-
cipital sutures are unfused.

"Juvenile": Individuals in which the molars are
fully erupted, the deciduous antemolar dentition
is erupted but still in the process of replacement
by the permanent teeth, and cranial sutures are
in the process of fusing. The eruption sequence
of the permanent teeth has been subdivided into
four stages by MacPhee (1987); these stages
have been accepted in this text unless otherwise
stated.

"Adult": Individuals with a fully erupted per-
manent dentition; cranial sutures are generally
fused, although their position is more or less
clearly marked.

Other abbreviations used are as follows: PN,
Pare National; RNI, Reserve Naturelle Integrate;
RS, Reserve Speciale; BM (NH), The Natural
History Museum, London (formerly British Mu-
seum [Natural History]); FMNH, Field Museum
of Natural History, Chicago; MCZ, Museum of
Comparative Zoology (Harvard), Cambridge;
MNHN, Museum National d'Histoire Naturelle,
Paris; SMG, Field catalog of S. M. Goodman;
UA, Departement de Biologie Animale, Univer-
site d' Antananarivo, Antananarivo; UMMZ, Uni-
versity of Michigan, Museum of Zoology, Ann
Arbor; and USNM, United States National Mu-
seum, Washington, D.C.

Systematics

Family Tenrecidae
Subfamily Tenrecinae

Setifer setosus Schreber, 1778

Holotype Unknown.

Type Locality Madagascar.

Key Features (see Key 1, p. 228) Dorsum
covered with spines; very short spinous tail pre-
sent. Skull moderately robust, dorsal profile

GOODMAN & JENKINS: LIPOTYPHLA

205

curved in lateral view; rostrum deep and broad;
interorbital region broad and elongated, frontals
posterodorsally inflated; interorbital region deeper
than braincase; braincase short, lambdoid crest
present. Dental formula 2/2 1/1 3/3 3/3 = 36; II
well developed, slightly shorter than C; short di-
astemata on either side of C and first lower pre-
molar (p2).

Remarks This species was not captured dur-
ing the 1996 survey of the PN de Marojejy. It was
observed on two occasions in the 450 m zone and
once at 775 m. It has been previously reported as
occurring in the reserve (Nicoll & Langrand.
1989). Setifer setosus was also found at low ele-
vation in the RS d'Anjanaharibe-Sud (Goodman
& Jenkins. 1998). This species occurs across the
complete length of the eastern humid forest, gen-
erally at lower-lying elevations, from the PN de
la Montagne d'Ambre in the north to the PN
d'Andohahela in the extreme south (Raxworthy &
Nussbaum, 1994; Goodman et al.. 1996b. 1999a).
It is also known from a variety of sites in western
deciduous forest (Ganzhorn et al.. 1996) and
spiny forest (Nicoll & Langrand. 1989).

Tenrec ecaudatus (Schreber, 1778)

Holotype Unknown.

Type Locality Unknown.

Key Features (see Key 1 . p. 228) The largest
of the Tenrecinae. Adult with dorsal pelage of
coarse bristly hair intermixed with soft spines;
short tail present. Skull elongated; rostrum with
deep sockets in ventrolateral region of the pre-
maxillae. which accommodate the lower canines
when the jaw is closed; interorbital region narrow,
elongated; braincase short, narrow, angular, pro-
nounced sagittal and lambdoid crests form deep
posterodorsal flanges. Dental formula 2/3 1/1 3/3
3/3 = 38; upper and lower canines very long, ro-
bust, and prominent; pronounced diastemata on
either side of C and posterior to C; short diaste-
mata posterior to first upper and lower premolars.

Remarks Tenrec ecaudatus was observed on
three occasions in the 450 m zone and once in the
775 m zone. Generally this species was noted in
the late afternoon or toward sunset, scuffling
along on the forest floor and presumably foraging
for food. It has been reported previously as oc-
curring in the reserve (Griveaud. 1960; Nicoll &
Langrand. 1989). In the RS d'Anjanaharibe-Sud.
this species was observed in the elevational zone
between 875 and 1260 m (Goodman & Jenkins,

1998). It is one of the most broadly distributed
tenrecids on Madagascar and occurs in a wide va-
riety of habitat types, including humid and decid-
uous forests as well as spiny bush.

Subfamily Oryzorictinae

Xticrogale brevicaudata G. Grandidier, 1899

Microgale breviceps Kaudern. 1918
Paramicrogale occidentalis Grandidier and
Petit. 1931

Lectotype MNHN 1986-387. adult male,
skull and skeleton, no skin, collector unknown.
Lectotype designation by MacPhee (1987, p. 6).

Type Locality Environs of Mahanara, 75
km S of Vohemar. NE coast of Madagascar.
(MacPhee [1987] provides coordinates of
1358'S. 4958'E for Antsirabe Avaratra. where
the Mahanara River crosses Route Nationale 5A.
Province d'Antsiranana.)

Referred Material FMNH 159652, 159653.
159654. and UA-SMG 8267: tributary of the
Manantenina River, 8 km NW of Manantenina.
Province d'Antsiranana. PN de Marojejy. 450 m.
1426.2'S, 4946.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) The following description is based on the
lectotype and two adult specimens from PN de
Marojejy. Body small to medium in size, tail short
relative to body length (TL as a percentage of HB
51.1. 51.7 in adult specimens from PN de Maro-
jejy [50% in the lectotype, based on measure-
ments in the original description]; ratio of TL to
CIL 1.76. 1.85 [1.52 in the lectotype]). Dorsal
pelage moderately short and coarse in texture,
brown with buffy brown speckling; ventral pelage
pale grayish brown, demarcated from dorsal col-
oration; tail and feet brown; tail scale hairs mod-
erately short, scales visible. Skull with short,
broad, and deep rostrum and moderately broad in-
terorbital region: braincase short, scarcely broader
than zygomatic process of maxillae; superior ar-
ticular facets angular and visible in dorsal view,
lambdoid crest prominent: mandible moderately
robust, corpus short and deep, coronoid process
broad. Dentition with short diastemata separating
II from 12 and 13 from C; 12 and 13 closely ad-
pressed: II one-third to one-half as tall as 12: C
prominent, accessory cusps poorly developed; tal-
onid of m3 well developed, hypoconid. hypocon-
ulid and entoconid ridge well marked, entoconid
poorly defined.

206

FTELDIANA: ZOOLOGY

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GOODMAN & JENKINS: LIPOTYPHLA

207

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Measurements External and cranial measure-
ments are presented in Table 11-1.

Reproduction The testes of the two adult
males (FMNH 159654 and 159652) collected
measured 8X5 and 9X8 mm, respectively,
whereas those of the juvenile (FMNH 159653)
measured 5X4 mm; nonetheless, the epididy-
mides of all three specimens were convoluted. No
females were collected during the survey.

Remarks M. brevicaudata was captured only
during the inventory of the PN de Marojejy in the
lowland forest at 450 m (Table 11-2). The modern
distribution of this species along the western coast
includes the deciduous forest of Kirindy and near
Maintirano and the slightly more humid habitat of
the RS de Manongarivo (MacPhee, 1987; Nicoll
& Langrand, 1989; Nicoll & Rathbun, 1990; Gan-
zhorn et al., 1996; Fig. 11-2). The details of the
Manongarivo record are not clear, but presumably
this species was found in the lowland forest,
which contains dry deciduous floristic elements
(Humbert, 1951). M. brevicaudata has recently
been collected in deciduous forest of the RN1
d'Ankarafantsika, between 160 and 200 m (Ran-
drianjafy Rasoloarisoa, 1993; Rakotondravony,
Randrianjafy Rasoloarisoa, and Goodman, un-
publ. data). In the extreme north, it is known from
the PN de la Montagne d'Ambre, in the ecotone
between dry and humid forest (Raxworthy &
Nussbaum, 1994; Goodman et al., 1996b). As
mentioned earlier, the lectotype of M. brevicau-
data comes from the Mahanara River region, be-
tween Vohemar (Iharana) and Sambava, approxi-
mately 50 km NE of the site where this species
was captured in the PN de Marojejy. There is no
evidence that M. brevicaudata occurs in the Per-
inet region, and the damaged specimen cited by
Eisenberg and Gould (1970, p. 30) as a record of
it at this site (USNM 341694) is a misidentified
M. cowani. In summary, this species occurs in a
variety of habitat types, from lowland humid for-
est, including the lower slopes of several moun-
tains, to dry deciduous forest. It seems to ring the
coastal regions of the island, from the northeast
near Sambava and Vohemar, north to the Antsi-
ranana region, and south to at least near Moron-
dava.

Further evidence of the broad habitat tolerance
of M. brevicaudata or at least of their apparent
former larger geographical range comes from
bones recovered from various Holocene deposits
on the island. Material of this species has been
recovered in the cave Lakoton'i Akanga, north of
Antsiranana, which may have been a former car-

208

FIELDIANA: ZOOLOGY

Table 11-2. Elevational distribution of tenrecids on the slopes of the PN de Marojejy.

Individuals captured at elevation (m)

Species

450

775

1225

1625

1875

(+)

(+)

(+)

(+)

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+
+

+
+
+

+

+

+

+

+

+

+

6

4

8

9

4

3

2

8

8

4

25

17

67

67

33

Setifer setosus
Tenrec ecaudatus
Microgale brevicaudata
Microgale cowani
Microgale dobsoni
Microgale fotsifotsy
Microgale gracilis
Microgale gymnorhyncha
Microgale longicaudata
Microgale monticola
Microgale parvula
Microgale principula
Microgale soricoides
Microgale talazaci
Oryzorictes hova

Total number of Lipotyphla

Total number of Microgale spp.

Microgale spp. (N = 12) in each zone (%)

Note Includes sight records (in parentheses) and information of species obtained in live traps and pitfall traps
from the 1996 survey.

44 U E

46 U E

48E

50 U E

Fig. 11-2. Sites where Microgale brevicaudata has been collected or reported from.

GOODMAN & JENKINS: LIPOTYPHLA

209

nivore den (MacPhee, 1987; Dewar & Rakoto-
vololona, 1992). A toothless mandible found at a
site in the extreme southwest has been tentatively
assigned to M. brevicaudata (MacPhee, 1986).
Further, two mandibles recovered from a cave
near Mahajanga were described tentatively as a
new species, M. breviceps (Kaudern, 1918),
which was subsequently synonymized with M.
brevicaudata (MacPhee, 1987).

Microgale cowani Thomas, 1882

Holotype BM (NH) 82.3.1.25, adult female,
body preserved in alcohol, skull extracted, col-
lected mid-February to mid-March 1880 by the
Reverend W. Deans Cowan.

Type Locality Ankafana Forest, eastern Bet-
sileo. (Ankafana = Ankafina, Fianarantsoa, Fian-
arantsoa Province, 2112'S, 4712'E; see Mac-
Phee [1987], Carleton & Schmidt [1990].)

Referred Material FMNH 159516, 159518,
159519, 159520, 159521, 159522, 159523,
159524, 159525, and 159655, and UA-SMG 8365
and 8381: Antranohofa, 11 km NW of Mananten-
ina, Province d'Antsiranana, PN de Marojejy,
1225 m, 1426.02'S, 4944.5'E; FMNH 159526,
159527, 159528, 159529, 159530, 159531,
159532, 159533, 159534, 159535, 159536,
159656, 159657, 159658, and 159659: tributary
at head of Andranomifototra River, 10.5 km NW
of Manantenina, Province d'Antsiranana, PN de
Marojejy, 1625 m, 1426.04'S, 4944.5'E; FMNH
159651, 159660, and 159661: source of Andran-
omifototra River, 11 km NW of Manantenina,
Province d'Antsiranana, PN de Marojejy, 1875 m,
1426.08'S, 4944.1'E.

Key Features (see Table 11-1 and Key 2, p.
228) Medium sized, tail moderately short, short-
er or subequal to HB. Pelage dark brown dorsally,
gray ventrally with a reddish brown wash; tail bi-
colored, dark brown dorsally, sharply demarcated
from paler reddish buff venter. Skull medium in
size, rostrum elongated. Dentition with pro-
nounced diastemata separating upper teeth from
II to P3, also lower teeth from i3 to p2. All ele-
ments of talonid of m3 present, including hypo-
conid, entoconid ridge, talonid basin, and ento-
conid.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
The ratio of males to females was 1:1.3 and that
of infants to adults was 1 : 8; no juveniles of this

species were collected during the survey. Two of
the adult females (FMNH 159660 and 159661)
were pregnant, each with three embryos (with one
embryo in the left and two in the right oviduct
[FMNH 159660] and with two embryos in the left
and one in the right oviduct [FMNH 159661]); the
embryos measured 21 mm and 13 mm, crown-to-
rump lengths, respectively. Two other females
(FMNH 159529 and 159651) were lactating, the
latter showing two placental scars in the left and
one in the right oviduct. Mammary formula: 1-0-
2 (N = 4), 0-1-2 (N = 5). The testes of three
adult males with convoluted epididymides
(FMNH 159655, 159657, and 159659) measured
10 X 7, 8 X 6, and 7X5 mm, respectively,
whereas those of the infant male (FMNH 159658)
with nonconvoluted epididymides measured 2 X
2 mm.

Remarks One specimen (FMNH 159651), al-
though readily assigned to this species on the ba-
sis of skull characters, is aberrant in that its tail
is longer than its head and body length.

M. cowani was previously reported from the
PN de Marojejy (Nicoll & Langrand, 1989; Duck-
worth, 1990). During the 1996 survey, it was cap-
tured between 1225 and 1875 m, which is very
similar to its elevational range (1260-1950 m) in
the RS d'Anjanaharibe-Sud (Table 11-3; Good-
man & Jenkins, 1998). These two sites are the
northernmost localities on the island for this wide-
spread species, which occurs in humid forests
along the Central High Plateau and eastern es-
carpment south to the PN d'Andohahela (Good-
man et al., 1996c; Goodman et al., 1998a, 1999a).

In the 1875 m zone, one pitfall line was placed
on a ridge, above forest line, in a zone of open
savannah. The only lipotyphlan captured in this
pitfall line was a single individual of M. cowani
(Table 1 1 -4), which supports evidence from other
localities that this species is not strictly forest-
dwelling (Langrand & Goodman, 1997).

Microgale dobsoni Thomas, 1884

Nesogale dobsoni Thomas, 1918

Holotype BM (NH) 84.10.20.1, immature
male, preserved in alcohol, skull extracted. Col-
lected February or March 1884 by W. Waters.

Type Locality Nandesen forest, Central Bet-
sileo (Nandihizana, 10 miles S of Ambusitra
manuscript note in Thomas' private copy of orig-
inal description, archived in BM [NH]. Nandihi-
zana, approximately 20 miles (30 km) SSW of

210

FIELDIANA: ZOOLOGY

Table 1 1-3. Elevational distribution of various Lipotyphla on mountains in eastern humid forests of Madagascar
(all sites sampled with similar trapping techniques).

Species

PN de la RS

Montagne PN de d'Anjanaharibe- PN PN

d'Ambre 1 Marojejy 2 Sud 2 d'Andringitra 4 d'Andohahela 5

(340-1350 m; (450-1875 m; (875-1950 m; (720-2450 m; (440-1875 m;

12S) 14S) 15S) 22S) 25S)

Tenrecidae

Hemicentetes nigriceps

1,350-2,050

Setifer setosus

650-

-660

450-775

875

810

440

Tenrec ecaudatus

650-

-1,350

450-775

875-1,260

720-810

440-1,200

Microgale brevicaudata

300-

-670

450

Microgale cowani

1,225-1,875

1,260-1,950

810-2,450

1,200-1,875

Microgale dobsoni

1,225-1,625

1,260

1,210-2,050

440-1,875

Microgale drouhardi

980-

-1,380

720-810

Microgale fotsifotsy

650-

-1,250

1,225

1,210-1,990

440-1,500

Microgale gracilis

1,875

1,210

1,500-1,875

Microgale gymnorhyncha

1,225-1,625

1,260-1,550

1,210-1,990

1,500-1,875

Microgale longicaudata

650-

-1,380

1,225

1,260-1,950

720-1.990

440-1,875

Microgale monticola

1,625-1,875

1,550-1,950

Microgale parvula

1,125-

-1,320

450-1,625

1,260-1,950

720-1,990

440-1,875

Microgale principula

1,625-1,875

875

Microgale soricoides

1,225-1,625

1,260-1,550

1,210-1,990

440-1,200

Microgale taiva

720-1,990

810-1,875

Microgale talazaci

660-

-1,380

450-1,625

1,260-1,550

1,990

Microgale thomasi

Oryzorictes hova

450-1,625

1,200-1,875

440-1,200

Oryzorictes tetradactylus

2,450

Soricidae

Suncus murinus 6

1,000-

-1,320

Total number of Tenrecidae

8

15

ll 7

16

12

Total number of Microgale spp.

6

12

9 8

11

10

1 Raxworthy and Nussbaum (1994); Goodman et al. (1996b); Jenkins et al. (1997).

2 This study.

3 Goodman and Jenkins (1998).

4 Goodman et al. (1996a); Langrand and Goodman (1997); S. M. Goodman (unpubl. data).

5 Goodman et al. (1999a).

6 Introduced to Madagascar.

7 An unidentified species of Hemicentetes, Oryzorictes talpoides (= hova), Microgale taiva, M. fotsifotsy, and M.
dryas were recorded on the slopes of the Anjanaharibe-Sud Massif during other surveys (Goodman & Jenkins, 1998).
Thus, with these additions, the number of Tenrecidae on the mountain would increase to 16 species.

8 With the information presented in the above footnote, this figure would be increased to 1 2 species.

Ambositra [see MacPhee, 1987]. Estimated as
2050'S, 4710'E).

Referred Material FMNH 159538, 159539,
159540, 159541, 159542, 159543, and 159662,
and UA-SMG 8352: Antranohofa, 11 km NW of
Manantenina, Province d'Antsiranana, PN de Ma-
rojejy, 1225 m, 1426.02'S, 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Large, TL subequal to or longer than HB.
Dorsal pelage brown, venter gray with buff wash.
Skull large and robust, sutures fused and obscure;
rostrum moderately broad, interorbital region
long; braincase angular, superior articular facets
very prominent, supraoccipital crests well devel-

oped; occipital region reduced. Diastemata be-
tween II and 12 and between 13 and C. The II
larger than 12, and i2 considerably larger than ca-
nine. Talonid of m3 reduced, hypoconid low, hy-
poconulid prominent, entoconid ridge and talonid
basin poorly defined, entoconid absent.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
Only adults were collected during the survey. Sex
ratio 1 female : 4 males. The testes of one male
(FMNH 159662) with convoluted epididymides
measured 7X3 mm. The mammary formula of
the single female (FMNH 159540) was 1-0-2.

GOODMAN & JENKINS: LIPOTYPHLA

211

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212

FIELDIANA: ZOOLOGY

Remarks In the PN de Marojejy, M. dobsoni
was captured at 1225-1625 m (Table 11-2). In the
RS d'Anjanaharibe-Sud, it was captured only at
1260 m (Table 1 1-3). This species is broadly dis-
tributed across much of the Central High Plateau
and eastern humid forest, from the mountains
skirting the Andapa Basin south to the PN
d'Andohahela (Goodman et al., 1996a, 1996c,
1998a; Goodman & Jenkins, 1998). On the An-
dohahela Massif, M. dobsoni was collected in the
elevational zone between 440 and 1 875 m, and on
the Andringitra Massif, from approximately 1210
m to above tree line, at 2050 m (Table 1 1-3).

Microgale fotsifotsy Jenkins, Raxworthy, and
Nussbaum, 1997

Microgale sp. A Jenkins, Goodman, and

Raxworthy, 1996

Microgale nov. sp. Goodman,

Andrianarimisa, Olson, and Soarimalala,

1996b

Holotype UMMZ 168468, adult male, pre-
served in alcohol, skull extracted, collected 13
January 1992 by Christopher J. Raxworthy.

Type Locality Antomboka River Fitsahana,
PN de la Montagne d'Ambre, Antsiranana Fivon-
dronana, Antsiranana Province, 1229'S, 4910'E,
altitude 650 m.

Referred Material FMNH 159663: Antra-
nohofa, 11 km NW of Manantenina, Province
d' Antsiranana, PN de Marojejy, 1225 m,
1426.02'S, 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Dorsal pelage pale grayish brown, soft in
texture, venter with lighter buffy wash; digits of
fore and hind feet and extreme tail tip contrast-
ingly lighter colored; pinnae prominent and con-
spicuous; fifth digit of hind foot elongated, scarce-
ly shorter than second. Skull with maxillary pro-
cess of zygoma at right angles to long axis of
cranium; braincase broad and short. 13 and i3
small, 13 slightly greater in crown height than dis-
tostyle of 12, i3 subequal in height to posterior
accessory cusp of i2; i2 greater in breadth than
il; C and c greater in crown height than P3 and
p3, respectively.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
The testes of the adult male measured 6X5 mm
and epididymides were convoluted.

Remarks Microgale fotsifotsy is broadly dis-

tributed along the complete length of the eastern
humid forests of Madagascar, from the PN de la
Montagne d'Ambre in the north to the PN
d'Andohahela in the extreme south (Jenkins et al.,
1997; Goodman et al., 1999a). It is currently un-
known from isolated forest fragments on the Cen-
tral High Plateau.

At some of the various sites where M. fotsifotsy
has been documented, it is captured infrequently.
The basis of this apparent rarity or absence at
some localities is unclear; it may simply be trap
shyness or low population density, although there
is some evidence of population cycling. In the PN
de la Montagne d'Ambre, pitfall lines were estab-
lished at identical sites during the same season
over 3 consecutive years (Goodman et al., 1997).
With standardized pitfall trapping efforts, two in-
dividuals were captured in 1994, eight in 1995,
and five in 1996.

In the PN de Marojejy, the single individual
captured during the 1996 survey was in the 1225
m zone. During the 1994 survey of the eastern
slopes of the RS d'Anjanaharibe-Sud, M. fotsifot-
sy was not recorded. However, this species was
captured with pitfall traps during a field trip con-
ducted by F Andreone in February 1996 on the
western slopes of Anjanaharibe-Sud (Goodman &
Jenkins, 1998).

Microgale gracilis Major, 1896

Oryzoryctes [sic] gracilis Major, 1896
Leptogale gracilis Thomas, 1918

Holotype BM (NH) 97.9.1.78, adult of un-
determined sex, skin and skull. Collected Novem-
ber 1894 by C. I. Forsyth Major.

Type Locality Ambohimitombo forest. (Am-
bohimitombo town, 43 km [by road] SE of Am-
bositra, 10 km into eastern forest; Fianarantsoa,
Fianarantsoa; 2043'S, 4726'E [see MacPhee,
1987]. MacPhee gives the altitude for this locality
variously as 1300 m [p. 6] and 1200 m [his Table
5], but, as pointed out by Carleton and Schmidt
[1990], the altitude recorded for this locality by
Major [1897] is higher [1500-1600 m].)

Referred Material FMNH 159664: source
of Andranomifototra River, 1 1 km NW of Man-
antenina, Province d'Antsiranana, PN de Maro-
jejy, 1987 m, 1426.08'S, 4944.1'E.

Key Features (see Table 11-1 and Key 2, pp.
228) Size large, TL shorter than HB. Pelage
dark brown dorsally with buff speckling. Muzzle
very long; large, naked rhinarium anteriorly retic-

GOODMAN & JENKINS: LIPOTYPHLA

213

ulated, striae on posterior region incomplete. Eyes
very small; ears small, partially concealed by pel-
age. Forefeet broad, claws enlarged. Skull very
elongated and gracile; rostrum slender, markedly
attenuated; braincase rounded, moderately broad
and long. Dentition reduced; upper incisors sub-
equal in height, incisors and canine very slender;
large diastemata between all anterior teeth, partic-
ularly P2 and P3; talons on molars very reduced,
resembling cingula; talonid of m3 slightly re-
duced, entoconid absent.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
The single adult female had one placental scar in
each oviduct; the mammary formula was 1-0-2.

Remarks This relatively poorly known and
difficult to capture shrew tenrec was obtained on
only one occasion in the PN de Marojejy. The
single individual was taken at 1875 m, in a Sher-
man trap placed on the ground along a passage-
way, in moss-covered roots and branches in a
transitional zone between mossy and sclerophyl-
lous forest. This species was not recorded in the
RS d'Anjanaharibe-Sud and is known only from
scattered localities on the Central High Plateau
and on the slopes of Andringitra and Andohahela
(MacPhee, 1987; Goodman et al., 1996a, 1996c,
1999a; Goodman & Jenkins, 1998). The record
reported for this species in the Fanovana area by
MacPhee (1987) is referable to M. gymnorhyncha
(Jenkins et al., 1996). On the basis of these com-
bined records, M. gracilis is known to have an
elevational distribution of 1210-2000 m.

Microgale gymnorhyncha Jenkins, Goodman,
and Raxworthy, 1996

Microgale gracilis (Major): MacPhee, 1987,
in part

Holotype FMNH 151807, adult female, pre-
served in alcohol, skull extracted (field number
SMG 6697), collected 13 December 1993 by
Steven M. Goodman and Christopher J. Raxwor-
thy.

Type Locality 38 km S of Ambalavao, RNI
d' Andringitra, on ridge E of Volotsangana River,
Fianarantsoa Province, 22H'39"S, 4658'16"E,
altitude 1625 m.

Referred Material FMNH 159544, 159665,
and 159666, and UA-SMG 8414: Antranohofa, 11
km NW of Manantenina, Province d' Antsiranana,
PN de Marojejy, 1225 m, 1426.02'S, 4944.5'E;
FMNH 159545: tributary at head of Andranomi-

fototra River, 10.5 km NW of Manantenina, Prov-
ince d' Antsiranana, PN de Marojejy, 1625 m,
1426.04'S, 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Large, TL shorter than HB. Dorsal pelage
dark brown, dark gray brown ventrally. Muzzle
very long, forming a proboscis; rhinarium very
large, with transversely striated naked region.
Eyes very small. Ears small, virtually concealed
in pelage. Forefeet broad, claws enlarged. Skull
long, moderately gracile; rostrum slender and
elongated; braincase short and broad. Dentition
moderately reduced, with long diastemata be-
tween all anterior teeth from II to P3 and i2 to
p3; talonid of m3 slightly reduced; talonid basin,
hypoconid, hypoconulid, and entoconid ridge pre-
sent, entoconid indicated.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
Sex ratio 1 male : 2 females; age ratio 1 juvenile :
3 adults. The mammary formula in both females
was 0-1-2.

Remarks The elevational zones where M.
gymnorhyncha was captured in the PN de Maro-
jejy and the RS d'Anjanaharibe-Sud are similar:
1225-1625 m in the former and 1260-1550 m in
the latter (Goodman & Jenkins, 1998). This spe-
cies is known to have a broad distribution from
the mountains surrounding the Andapa Basin
south to the PN d' Andohahela (Goodman et al.,
1999a). It has been collected at sites on the Cen-
tral High Plateau and along the eastern escarp-
ment (Jenkins et al., 1996; Goodman et al., 1998a,
1998b).

The elongated rostrum and rhinarium, reduced
dentition, small eyes and ears, digging claws,
short forelimbs, and dense short pelage of M.
gymnorhyncha are very reminiscent of M. graci-
lis, and these two species probably occupy a sim-
ilar ecological niche. They are known to occur in
sympatry in the PN d'Andohahela and PN
d' Andringitra (Jenkins et al., 1996; Goodman et
al., 1996a, 1999a), where they were captured in
the same elevational zone and, in a few cases, in
the same pitfall line. Whether there is overlap in
the prey taken by these two species would be an
interesting line of research.

Microgale longicaudata Thomas, 1882

Microgale majori Thomas, 1918: MacPhee,
1987

Holotype BM(NH) 82.3.1.15, adult female,
body preserved in alcohol, skull extracted, col-

214

FIELDIANA: ZOOLOGY

lected mid-February to mid-March 1880 by the
Reverend W. Deans Cowan.

Type Locality Ankafana Forest, eastern Bet-
sileo. (Ankafana = Ankafina, Fianarantsoa, Fian-
arantsoa Province; 2112'S, 4712'E; see Mac-
Phee, 1987; Carleton & Schmidt, 1990).

Referred Material FMNH 159546, 159667:
Antranohofa, 1 1 km NW of Manantenina, Prov-
ince d'Antsiranana, PN de Marojejy, 1225 m,
1426.02'S, 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Small in size; tail very long, more than
twice as long as HB; distal portion of tail naked
and transversely wrinkled on dorsal surface; fifth
hind digit elongated, subequal in length to second
digit. Dorsal pelage reddish brown, venter gray
with bright reddish buff or buff wash. Skull small,
rostrum moderately short; braincase moderately
narrow and long. Diastemata present between II
and 12 and either side of C and P2; well-devel-
oped anterior and posterior accessory cusps pre-
sent on 12, C, and P2; C subequal to or taller than
II; P4 scarcely greater in crown height than P3.
The p2 caniniform. Talonid of m3 with low hy-
poconid, hypoconulid well developed, narrow tal-
onid basin, reduced entoconid ridge, and entoco-
nid absent.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
Sex ratio 1:1. Mammary formula 1-0-2 (N = 1).

Remarks This species is one of the more
widespread shrew tenrecs. It is known from the
complete breadth of the eastern humid forest
(Montagne d'Ambre to Andohahela) and from
several sites on the Central High Plateau (Mac-
Phee, 1987; Raxworthy & Nussbaum, 1994;
Goodman et al., 1996c, 1998a, 1998b, 1999a). A
single specimen collected in the Kirindy Forest,
near Morondava, has been tentatively identified as
part of the M. longicaudata cluster (Ade, 1996).

M. longicaudata was captured only twice dur-
ing the inventory of the PN de Marojejy, and only
in the 1225 m zone. In the RS d'Anjanaharibe-
Sud, with nearly the same pitfall trapping effort,
it was trapped twice in the 1260 m zone, was not
recorded in the 1550 m zone, and was obtained
twice in the 1950 m zone. Detailed information
on the elevational range of M. longicaudata is dif-
ficult to glean from pitfall traps, since capture is
infrequent. The elevational distribution of this
species with regard to M. principula, a morpho-
logically similar form, is discussed below, in the
account of the latter species.

Microgale monticola Goodman and Jenkins,
1998

Holotype FMNH 154012 (field number
SMG 7020), adult female, skin and skull, collect-
ed 15 November 1994 by Steven M. Goodman.

Type Locality 11 km WSW of Befmgitra,
RS d'Anjanaharibe-Sud, 1444'S, 4926'E, alti-
tude 1550 m.

Referred Material FMNH 159549, 159550,
159551, 159672, and 159673: Antranohofa, 11
km NW of Manantenina, Province d'Antsiranana,
PN de Marojejy, 1225 m, 1426.02'S, 4944.5'E;
FMNH 159517, 159552, 159553, 159554,
159555, 159556, 159674, 159675, and 159676,
and UA-SMG 8486: tributary at head of Andran-
omifototra River, 10.5 km NW of Manantenina,
Province d'Antsiranana, PN de Marojejy, 1625 m,
1426.04'S, 4944.5'E; FMNH 159557, 159677,
159678, and 159679, and UA-SMG 8500: source
of Andranomifototra River, 1 1 km NW of Man-
antenina, Province d'Antsiranana, PN de Maro-
jejy, 1875 m, 1426.08'S, 4944.1'E.

Key Features (see Table 11-1 and Key 2, p.
228) Medium-sized Microgale with tail longer
than head and body. Pelage dark dorsally and ven-
trally. Skull moderately robust, with broad inter-
orbital region and broad, rounded braincase. Up-
per and lower canines robust, P2 and p2 large, P2
with well-developed anterior and posterior acces-
sory cusps.

Measurements External and cranial measure-
ments are given in Table 11-1.

Population Structure and Reproduction
Only adults were collected during the survey, and
the sex ratio of females to males was 1 : 1.7. Of
the females, one (FMNH 159676) was pregnant,
another (FMNH 159553) lactating, and a third
(FMNH 159554) pregnant and lactating. Both
pregnant specimens contained single embryos in
both left and right oviducts; in FMNH 159554,
which appeared close to parturition, the well-de-
veloped embryo measured 30 mm (crown-to-
rump length), whereas the small embryo in
FMNH 1 59676 measured only 3 mm. Another fe-
male (FMNH 159675) had single placental scars
in both oviducts. Mammary formula: 1-0-2 (N =
5), 0-1-2 (N = 1). Four of the adult males had
convoluted epididymides and testes measuring 4
X 3 mm (FMNH 159673), 5 X 5 mm (FMNH
159674 and 159678), and 6 X 5 mm (FMNH
159672); in a fifth male with fully erupted, worn,
permanent dentition (FMNH 159679) but with

GOODMAN & JENKINS: LIPOTYPHLA

215

nonconvoluted epididymides, the testes measured
4X3 mm.

Remarks This recently named species is cur-
rently known only from the mountains surround-
ing the Andapa Basin. In the PN de Marojejy, it
was recorded in the elevational zone between
1625 and 1875 m. and in the RS d'Anjanaharibe-
Sud, from 1550 to 1950 m (Tables 11-2. 11-3).
At both of these sites, it was one of the most
frequently captured shrew tenrecs. The morpho-
logically similar M. thomasi is not known to occur
in sympatry with M. monticola. M. thomasi has
been documented at a variety of sites on the Cen-
tral High Plateau and along the eastern escarp-
ment, and its known range is from the Andrano-
may Forest south to the PN d'Andohahela (Good-
man et al., 1996c. 1988a, 1999a).

Microgale parvula G. Grandidier, 1934

Microgale pulla Jenkins. 1988

Holotype MCZ 45465, juvenile male, body
preserved in alcohol, skull extracted, collected by
M. Drouhard.

Type Locality environs of Diego-Suarez
(Antsiranana, approximately 1216'S. 4918'E;
see MacPhee [1987]; probably Montagne
d'Ambre; see Jenkins et al. [1997]).

Referred Material FMNH 159558. 159559.
159680. and 159681: tributary of the Mananteni-
na River. 8 km NW of Manantenina. Province
d" Antsiranana. PN de Marojejy. 450 m. 1426.2'S.
4946.5'E: FMNH 159560. 159682. 159683. and
159684, and UA-SMG 8292: tributary of Man-
antenina River, 10 km NW of Manantanina, Prov-
ince d' Antsiranana. PN de Marojejy. 775 m,
1426.0'S, 4945.7'E; FMNH 159561. 159562.
159563. and 159685: Antranohofa. 11 km NW of
Manantenina. Province d' Antsiranana. PN de Ma-
rojejy. 1225 m. 1426.02'S, 4944.5'E; FMNH
159686: tributary at head of Andranomifototra
River. 10.5 km NW of Manantenina. Province
d' Antsiranana, PN de Marojejy, 1625 m.
1426.04'S. 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Very small, TL slightly shorter than HB.
Dorsal pelage dark brown, ventral pelage dark
gray brown, tail uniform dark gray. Skull very
small and delicate, rostrum slender, braincase
shallow and long, occipital condyles posterodor-
sally oriented. Diastemata between II and 12 and
on either side of C and P2: anterior and posterior
accessory cusps present on 12. 13. and P2. Diaste-

ma between c and p2. Talonid of m3 with well-
developed hypoconulid but with reduced hypo-
conid. entoconid. and entoconid ridge, and with
narrow, shallow talonid basin.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
Sex ratio 1:1; juvenile to adult ratio 1 : 2.3. One
female (FMNH 159682) had a single placental
scar in the left and two in the right oviduct. Mam-
mary formula: 0-1-2 (N - 1), 1-0-2 (N = 1). Two
adult males (FMNH 159681 and 159683) with
convoluted epididymides had testes measuring 5
X 3 mm and 3X3 mm. respectively; the testes
of a juvenile male with nonconvoluted epididy-
mides measured 2 X 1 mm.

Remarks Although never particularly com-
mon in any transect zone in the PN de Marojejy.
as indicated by pitfall trap captures, M. parxula
has a broad elevational range on the massif. It
occurs from lowland forest at 450 m to montane
forest at 1625 m (Table 11-2). In the RS
d'Anjanaharibe-Sud. this species was recorded
between 1260 and 1950 m (Table 11-3). Given
the total number of animals of this species cap-
tured in pitfall traps at these two sites, it may be
surmised that they have relatively similar eleva-
tional ranges on the two massifs.

When MacPhee (1987) conducted his revision
of Microgale, only a single specimen of M. par-
vula. the holotype. was available for study. Since
then, almost exclusively as a result of the use of
pitfall trapping techniques, this very small species
is now known from numerous sites along the
complete length of the eastern humid forest, from
the PN de la Montagne d'Ambre south to the PN
d'Andohahela (Raxworthy & Nussbaum, 1994;
Goodman et al., 1996a, 1996b, 1999a; Goodman
& Jenkins. 1998) and at several sites on the Cen-
tral High Plateau (Goodman et al.. 1998a, 1998b).
Throughout this zone, it has a broad elevational
range, from 440 to 1990 m.

Microgale principula Thomas, 1926

Microgale sorella Thomas. 1926: MacPhee,
1987

Holotype BM (NH) 25.8.3.15, adult female,
body preserved in alcohol, skull extracted, col-
lected by C. Lamberton.

Type Locality Midongy du Sud, SE Mada-
gascar (Midongy Atsimo. 2335'S, 4701'E; see
MacPhee [1987]).

216

FTELDIANA: ZOOLOGY

Referred Material FMNH 159547 and
159668, and UA-SMG 8489: tributary at head of
Andranomifototra River, 10.5 km NW of Man-
antenina, Province d'Antsiranana, PN de Maro-
jejy, 1625 m, 1426.04'S, 4944.5'E; FMNH
159548, 159669, 159670, and 159671: source of
Andranomifototra River, 1 1 km NW of Manan-
tenina, Province d'Antsiranana, PN de Marojejy,
1875 m, 1426.08', 4944.1'E.

Key Features (see Table 11-1 and Key 2, p.
228) Medium sized, tail very long, more than
twice as long as HB; distal portion of tail naked
and transversely wrinkled on dorsal surface; fifth
hind digit elongated, subequal in length to sec-
ond digit. Pelage distinctly bicolored, reddish
brown dorsally, gray with buff wash ventrally.
Skull medium in size, rostrum moderately short
and broad, braincase moderately narrow. Short
diastemata between II and 12 and on either side
of C and P2; 12 and 13 more or less in contact;
well-developed anterior and posterior accessory
cusps present on 12, C, and P2; II greater in
crown height than C; P4 distinctly greater in
crown height than P3; lower p2 moderately ca-
niniform, talonid of m3 with low hypoconid,
well-developed hypoconulid, broad talonid ba-
sin, reduced entoconid ridge, and entoconid ab-
sent.

Measurements External and cranial mea-
surements are presented in Table 11-1.

Population Structure and Reproduction
Only adults were collected during the survey.
Sex ratio 1:1. Mammary formula: 1-0-2 (N =
3). Two males (FMNH 159669 and 159671) with
permanent dentition but nonconvoluted epididy-
mides had testes measuring 4X3 mm and 6 X
4 mm, respectively.

Remarks It was previously proposed that the
geographical range of M. principula was con-
fined to the southern portion of the eastern humid
forest (MacPhee, 1987). The results of recent
surveys demonstrate that it has a much broader
distribution and occurs from at least the moun-
tains surrounding the Andapa Basin south to the
PN d'Andohahela (Goodman et al., 1999a). Cu-
riously, this species seems to have a patchy dis-
tribution. Intensive survey work in the PN
d'Andringitra failed to find it (Goodman et al.,
1996a), although it is known to occur not far to
the south, near Midongy du Sud and Ranomena
(MacPhee, 1987), and it is relatively common
less than 80 km to the northeast, in the PN de
Ranomafana (specimens in USNM).

Microgale soricoides Jenkins, 1993

Holotype BM (NH) 91.565, adult male pre-
served in alcohol, skull extracted. Collected 13
April 1991 by Christopher J. Raxworthy.

Type Locality Mantady National Park (PN
de Mantady), approximately 15 km north of Per-
inet (Andasibe), 1851'S, 4827'E, in primary rain
forest, altitude 1100-1150 m.

Referred Material FMNH 159564, 159565,
159566, 159568, 159687, 159688, and 159689,
and UA-SMG 8395 and 8484: Antranohofa, 11
km NW of Manantenina, Province d'Antsiranana,
PN de Marojejy, 1225 m, 1426.02'S, 4944.5'E;
FMNH 159567, 159568, 159569, 159570: tribu-
tary at head of Andranomifototra River, 10.5 km
NW of Manantenina, Province d'Antsiranana, PN
de Marojejy, 1625 m, 1426.04', 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Size large, TL subequal to or longer than
HB. Pelage light buff brown dorsally, gray-brown
ventrally with reddish buff wash. Skull moderate-
ly large and robust, rostrum and interorbital re-
gion broad, braincase short and broad; supraoc-
cipital ridge present. II markedly robust and pro-
odont. The il and i2 robust and procumbent, i2
smaller than il but larger than c. P2 and p2 very
small, with a single root. Talonid of m3 reduced
to very low hypoconid, an oblique crest, and a
prominent hypoconulid.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
Only adults were collected during the survey. The
ratio of males to females was 1 : 1.25. Two fe-
males (FMNH 159687 and 159688) were preg-
nant, the former with one embryo in the left and
two in the right oviduct, the embryos measuring
23 mm crown-to-rump length; the latter with an
embryo in each oviduct, 17 mm crown-to-rump
length. Mammary formula: 1-0-2 (N = 5). One
male (FMNH 159689) with convoluted epididy-
mides had testes measuring 6X4 mm.

Remarks On the basis of the 1996 inventory
of the PN de Marojejy, M. soricoides occurs in
the 1225 m and 1625 m zones (Table 11-2), where
it was relatively common; all individuals were
captured in pitfall traps. At other sites it is not
uncommon to capture this species in Sherman live
traps (e.g., Goodman et al., 1999a). The eleva-
tional range of M. soricoides in the RS
d'Anjanaharibe-Sud was 1260-1550 m (Good-
man & Jenkins, 1998), very similar to that on the
Marojejy Massif.

GOODMAN & JENKINS: LIPOTYPHLA

217

Just a few years after its description, we now
know that M. soricoides is a widely distributed
and relatively common species. Extensive surveys
conducted in the PN de la Montagne d'Ambre
have failed to find this species (Raxworthy &
Nussbaum, 1994; Goodman et al., 1996b), and the
known northern limit is along the slopes of An-
janaharibe-Sud and Marojejy. We strongly suspect
that this species will be found in the Sambirano,
particularly on the Manongarivo and Tsaratanana
Massifs. To the south of the Andapa Basin it is
broadly distributed in the eastern humid forest and
on the Central High Plateau to the southern limit
of this forest type in the PN d'Andohahela (Jen-
kins, 1993; Goodman & Jenkins, 1998; Goodman
et al., 1998a, 1999a). Further, this species occurs
across a wide elevational range: in the PN
d'Andohahela, it was recorded between 810 and
1875 m (Goodman et al., 1999a).

M. soricoides, with its massive upper incisors,
is clearly predatory. At numerous sites, this spe-
cies, as well as M. dobsoni and M. talazaci, has
been found trapped in the same pitfall bucket with
smaller species of Microgale, and more often than
not the M. soricoides has partially eaten the other
individual. In some cases, the only evidence that
remains of the consumed shrew tenrec is a portion
of tail or rostrum.

Microgale talazaci Major, 1896

Nesogale talazaci Thomas, 1918

Holotype BM (NH) 97.9.1.107, adult female;
skin, skull, and skeleton. Collected 22 May 1896
by C. I. Forsyth Major.

Type Locality Forest of the Independant
Tanala of Ikongo, in the neighborhood of Vinan-
itelo, one day's journey south of Fianarantsoa
(2144'S, 4716'E).

Referred Material FMNH 159690: tribu-
tary of the Manantenina River, 8 km NW of Man-
antenina, Province d'Antsiranana, PN de Maro-
jejy, 450 m, 1426.2'S, 4946.5'E; FMNH
159537, 159571, 159691, 159692, and 159693:
tributary of Manantenina River, 10 km NW of
Manantenina, Province d'Antsiranana, PN de Ma-
rojejy, 775 m, 1426.0'S, 4945.7'E; FMNH
159572, 159573, 159574, 159575, and 159576:
Antranohofa, 1 1 km NW of Manantenina, Prov-
ince d'Antsiranana, PN de Marojejy, 1225 m,
1426.02'S, 4944.5'E; FMNH 159694 and UA-

SMG 8450: tributary at head of Andranomifototra
River, 10.5 km NW of Manantenina, Province
d'Antsiranana, PN de Marojejy, 1625 m,
1426.04'S, 4944.5'E.

Key Features (see Table 11-1 and Key 2, p.
228) Very large, TL longer than HB. Dorsal pel-
age brown, venter gray with reddish buff wash.
Skull very large and robust, sutures fused and ob-
scure; rostrum broad, interorbital region long, par-
allel-sided; braincase angular, short relative to cra-
nial length, superior articular facets very promi-
nent, supraoccipital crests well developed, occip-
ital region very reduced, occipital condyles visible
in dorsal view. Small diastemata between II and
12 and between 13 and C. II larger than 12; lower
i2 considerably larger than canine. Talonid of m3
reduced, hypoconid low, hypoconulid well
marked, entoconid ridge and talonid basin poorly
defined, entoconid absent.

Measurements External and cranial measure-
ments are presented in Table 11-1.

Population Structure and Reproduction
The sex ratio of males to females was 1 : 2, the
juvenile to adult ratio 1 : 9. Mammary formula: 1-
0-2 (N = 1), 1-1-2 (N = 1). The testes of one
adult male (FMNH 159694), with convoluted ep-
ididymides, measured 8X6 mm.

Remarks M. talazaci, the largest species of
shrew tenrec, has a broad distribution across much
of the humid forest zones of the island, from the
Antsiranana region, including the PN de la Mon-
tagne d'Ambre (MacPhee, 1987; Goodman et al.,
1996b) south across the eastern escarpment and
several sites on the Central High Plateau, to at
least the Vondrozo region (MacPhee, 1987; Good-
man & Jenkins, 1998; Goodman et al., 1998a).
This species was not found in the PN
d'Andohahela (Goodman et al., 1998a). Its pres-
ence in the PN de Marojejy was reported previ-
ously by Nicoll and Langrand (1989) and by
Duckworth (1990), and it is also known to occur
in the RNI de Tsaratanana (Albignac, 1970).

In the PN de Marojejy, it was found to have an
elevational range of 450-1625 m (Table 11-2),
which is slightly broader than its range in the RS
d'Anjanaharibe-Sud, where it was documented at
1260-1550 m (Table 11-3). In the PN
d'Andringitra, this species' known altitudinal dis-
tribution is more restricted it was not found in
the zone from 720 to 1650 m but was trapped near
tree line, at 1990 m (Goodman et al., 1996a;
Goodman, unpubl. data).

218

FIELDIANA: ZOOLOGY

Oryzorictes hova A. Grandidier, 1870

Oryzorictes talpoides G. Grandidier and
Petit, 1930

Holotype MNHN CG 1887-874, adult fe-
male, preserved in alcohol, skull extracted, col-
lection date unknown.

Type Locality Ankaye et Antsianak (Ankaye
= Ankay, along the Mangoro River near lac Ala-
otra; Antsianak = Antsianaka, to the east of lac
Alaotra [Viette, 1991]).

Referred Material FMNH 159577 and
159695: tributary of the Manantenina River, 8 km
NW of Manantenina, Province d'Antsiranana, PN
de Marojejy, 450 m, 1426.2'S, 4946.5'E; FMNH
159578: tributary at head of Andranomifototra
River, 10.5 km NW of Manantenina, Province
d'Antsiranana, PN de Marojejy, 1625 m,
1426.04'S, 4944.5'E.

Key Features (see Key 1, pp. 228) Pelage
soft, slightly iridescent; TL approximately half
HB (TL:HB mean 49.51 2.70, range 46.0-
53.9; N =* 5); forefeet with very enlarged claws;
broad naked rhinarium; eyes very small; ears
small, concealed in pelage. Skull moderately ro-
bust, premaxillae dorsolateral^ flared, braincase
short, broad, and deep, lambdoid crests well-
marked; 13 very small, approximately as tall as
distostyle of 12; diastema present between 13 and
C, forming a sulcus to accommodate the distal tip
of c; C and c markedly taller than all other teeth;
distostyle of C very small.

Measurements External measurements are
presented in Table 11-1.

Population Structure and Reproduction
Only adults were collected during the survey. The
single female (FMNH 159577) was perforate, and
the mammary formula was 0-1-2. One of the
males (FMNH 159695) with convoluted epididy-
mides had testes measuring 9X8 mm.

Remarks This species is generally difficult to
trap, probably because of its semifossorial habits.
Captures of this animal in pitfall traps or standard
live traps is sporadic and seems to be more com-
mon after heavy rain, when individuals may be
more active on the ground surface. In the PN de
Marojejy, this species was obtained in the 450 and
1625 m zones (Table 11-2); it probably occurs
across this complete elevational range. Local
guides that joined the 1996 Marojejy survey noted
that this distinctive animal is common in the rice
fields near Mandena and Manantenina.

Oryzorictes hova is probably more common
and more broadly distributed than is currently

known. It has not been captured in the PN de la
Montagne d'Ambre (Rax worthy & Nussbaum,
1994; Goodman et al., 1996b). No individual of
this species was trapped on the 1994 mission to
the eastern slopes of the RS d'Anjanaharibe-Sud,
but in February 1996 it was collected by F An-
dreone on the western slopes of this massif
(Goodman & Jenkins, 1998). The northernmost
known locality for this species in the eastern hu-
mid forests appears to be the mountains surround-
ing the Andapa Basin. It is known from a variety
of forested and nonforested sites along the eastern
escarpment and Central High Plateau south to the
PN d'Andohahela (Goodman et al., 1998b,
1999a). It is also known from the Marovoay re-
gion, several sites on the Masoala Peninsula, and
the Nosy Mangabe (specimens in BM(NH)).

Discussion

General

A total of 1,155 pitfall bucket-days were ac-
crued during the inventory of the PN de Marojejy,
which was conducted between 3 October and 20
November 1996. This effort was divided between
five elevational zones: 264 bucket-days at 450 m,
264 days at 775 m, 231 days at 1225 m, 231 days
at 1625 m, and 165 days at 1875 m (Table 11-4).
The pitfall traps yielded 117 small mammals, in-
cluding 107 Microgale, 2 Oryzorictes, 1 Eliurus
majori, 2 E. minor, and 5 Voalavo gymnocaudus
(for rodents, see Chapter 12). Furthermore, 3,390
trap-nights, during which small-mammal traps
with a standard baiting regimen were used, were
also accrued during the inventory (Chapter 12),
and 10 (0.29%) lipotyphlans were captured. The
following species were obtained in standard mu-
seum traps: Microgale cowani, M. gracilis, M.
talazaci, and Oryzorictes hova. In three cases,
shrew tenrecs were captured in an elevational
transect by means of standard museum traps and
were not obtained by means of the pitfall devices
in the same zone: Microgale talazaci at 450 m,
M. gracilis at 1875 m, and Oryzorictes hova at
1625 m. Further, the specimen of M. gracilis cap-
tured in the 1875 m zone is our only record of
this species on the Marojejy Massif.

The combined trapping results, with pitfalls and
standard live traps, found 13 species of Tenreci-
dae (12 species of Microgale plus Oryzorictes
hova) in the PN de Marojejy. Further, individuals

GOODMAN & JENKINS: LIPOTYPHLA

219

fzoo-155-06-f03

of Tenrec ecaudatus and Setifer setosus were ob-
served on the massif but were not trapped. Thus,
in total, 15 species of Tenrecidae were found at
this site during the 1996 survey. The only lipo-
typhlans previously reported from the reserve but
not found during our inventory are Hemicentetes
semispinosus and Microgale pusilla (Nicoll &
Langrand, 1989; Duckworth, 1990). The absence
of these two animals in the results of the 1996
faunal inventory is discussed below (p. 221). Be-
fore we proceed with the analysis of the trapping
results, it is important to assess whether the sam-
pling effort extended in the PN de Marojejy was
sufficient to allow some confidence in our mea-
sure of lipotyphlan species richness within each
elevational zone.

Species Accumulation Curves and Confidence
in Measurement of Species Richness

The total number of species known from each
elevational zone was plotted against sampling ef-
fort (33 pitfall bucket-days per 24-hour period) to
determine whether the number of cumulative ten-
rec species at each site reached a plateau (Fig. 1 1-
3a). An examination of these curves shows that,
in the 450 and 1875 m zones and, to a slightly
lesser extent, in the 775 m zone, the accumulation
of previously unrecorded species was rapid. One
or two nights were sufficient to reach an apparent
plateau in the species of lipotyphlans readily cap-
tured by pitfalls (see below). Of the five sites sam-
pled, these three elevational sites had the lowest
species richness (Table 11-2). Thus, it is not sur-
prising that plateaus were reached relatively
quickly. At 1225 and 1625 m, the zones with the
greatest number of lipotyphlans species on the
slopes of Marojejy, clear plateaus were not
reached. In the former zone, a new species (M.
fotsifotsy) was added the fifth night of pitfall trap-
ping, and in the latter zone, an additional species
(M. dobsoni) was added on the sixth and final
night.

Information on the accumulation of new spe-
cies in each elevational zone, based on pitfall
traps, can be summarized as follows: in the 450
m zone, no new species was added after 66 pitfall
bucket-days (total, 3 species in 264 pitfall bucket-
days); in the 775 m zone, after 198 pitfall bucket-
days (total, 2 species in 264 pitfall bucket-days);
in the 1225 m zone, after 198 pitfall bucket-days
(total, 9 species in 231 pitfall bucket-days); in the
1625 m zone, after 198 pitfall bucket-days (total,

t r

132 165 198 231 264

Bucket-days

33 66 99 132 165 198 231 264

Bucket-days

Fig. 1 1-3. Species accumulation curves (A) and pit-
fall trap success (B) plotted for each elevational zone in
the PN de Marojejy against the total number of bucket-
nights. The pitfall lines were placed in five different ele-
vational zones (450, 775, 1225, 1625, and 1875 m). In-
formation from the three lines in each zone is combined.

8 species in 231 pitfall bucket-days); and in the
1875 m zone, after 33 pitfall bucket-days (total, 3
species in 165 pitfall bucket-days). The leveling
out of these curves within each elevational zone

220

FIELDIANA: ZOOLOGY

through time did not generally coincide with a
decline in pitfall trap success (Fig. 11 -3b), al-
though the number of animals captured in each
elevational zone was generally higher at the be-
ginning, rather than at the end, of each trapping
season.

The Microgale species that are most difficult to
capture in pitfall traps are the large-bodied ani-
mals, particularly M. talazaci and, to a lesser ex-
tent, M. dobsoni. Oryzorictes, with its apparent
subterranean mode of life, is also captured infre-
quently by this trapping system. In the 450 m
zone, not a single M. talazaci was captured in the
pitfall traps, but one was captured with a Sherman
live trap on the second night of trapping (see
Chapter 12). In the 775 m zone, this species was
captured in Sherman live traps on the first, sec-
ond, and fourth nights of trapping and was not
obtained by pitfall traps until the sixth night. At
1 625 m, an Oryzorictes hova was trapped the sev-
enth and final night of trapping, but it was not
captured at this elevation in the pitfall traps. Fi-
nally, at the 1 875 m site, the sole specimen of M.
gracilis captured during the inventory of the PN
de Marojejy was obtained with a Sherman trap.

In general, we feel that our combined trapping
results (pitfall traps with standard traps) provide
a good estimate of the actual species of lipotyph-
lans occurring within each elevational zone. (For
further discussion of this point, see Goodman &
Jenkins, 1998, p. 156.) Little previous information
on the small mammals of this site is available
(Griveaud, 1960; Nicoll & Langrand, 1989; Duck-
worth, 1990). Two species have been reported
from the PN de Marojejy that were not encoun-
tered during the 1996 inventory: Microgale pus-
ilia and Hemicentetes semispinosus. The former
species was collected at 800 m in the northwest-
ern portion of the reserve near the Antsahaber-
oakahely River (Duckworth, 1990). The specimen
deposited in the Pare Botanique et Zoologique de
Tsimbazaza, Antananarivo, could not be found in
April 1998, and it is impossible to verify the de-
termination. We suspect that the animal was mis-
identified and is probably referable to the very
similarly small-sized M. parvula. M. pusilla was
not recorded in the nearby RS d'Anjanaharibe-
Sud (Goodman & Jenkins, 1998), and the nearest
known locality for this species with respect to
Marojejy is in the Didy region, approximately 400
km to the south. Duckworth (1990) reported ob-
serving H. semispinosus on 2 September in the
southwestern portion of the reserve. An uniden-
tified species of Hemicentetes was observed by F

Andreone on the western slopes of RS
d'Anjanaharibe-Sud (Goodman & Jenkins, 1998).
H. nigriceps is known only from the central high
plateau and higher altitudes of the eastern escarp-
ment, and the animal observed in the RS
d'Anjanaharibe-Sud is almost certainly H. semis-
pinosus, as indicated by the geographical distri-
bution. This species was not recorded during an
inventory of this reserve between mid-October
and late December 1994, and it seems distinctly
less common in the warm season, before heavy
rains (September through December) than during
the heart of the rainy season (January through
April). It has been reported from the Masoala
Peninsula, near Rantabe, and the Mananara region
(Nicoll & Langrand, 1990; Stephenson, 1995).
There is little doubt that H. semispinosus occurs
in the PN de Marojejy, and it was not recorded
during the 1996 inventory of the reserve.

Virtually all of the species of tenrecs that might
be expected to occur in the reserve were found
during the 1996 survey. The only exceptions to
this are M. dryas and M. taiva. The former species
has been identified from owl pellets collected at
the edge of the RS d'Anjanaharibe-Sud but not
obtained during the inventory of that reserve in
1994, and the latter species was captured along
the western slopes of the Anjanaharibe-Sud Mas-
sif (Goodman & Jenkins, 1998).

Several other studies have shown that pitfall
buckets are an excellent method to document the
diversity of Malagasy lipotyphlans living in hu-
mid forests. The use of this technique, combined
with standard live traps, over the course of a rapid
faunal inventory probably gives an excellent, al-
though not 100% complete, estimate of the spe-
cies living in an area.

The utility of pitfall traps in augmenting cap-
ture rates of Microgale and subsequently provid-
ing material for systematic studies can be nicely
illustrated with M. gymnorhyncha and M. fotsi-
fotsy, both of which have only recently been de-
scribed. These two species are now known to be
widely distributed throughout much of the humid
forest on Madagascar. Both of these species had
been represented in museum collections before
their "discovery" during recent biological inven-
tories. This newly collected material was neces-
sary to clearly diagnose these new forms and to
establish species limits.

Elevational Associations Among Lipotyphlans

Elevational Distribution No oryzorictine
species was trapped in all of the five zones sam-

GOODMAN & JENKINS: LIPOTYPHLA

221

pled along the elevational transect of the PN de
Marojejy (Table 1 1-2). Microgale parvula and M.
talazaci, recorded at the four sites between 450
and 1625 m, appear to occupy the broadest alti-
tudinal band among the Microgale found within
the reserve. For the former species, this finding is
in accordance with previous results from various
surveys (Table 11-3), where it occurs from low-
land to montane habitats. The case for M. talazaci
is somewhat different. At other surveyed sites,
this species apparently does not have as broad an
elevational range as in the PN de Marojejy (Table
1 1-3). This may in part be an artifact of the num-
ber of individuals captured at these other sites.
Only one species, M. brevicaudata, was restricted
to the lowlands of the PN de Marojejy. Several
species are apparently limited to the middle ele-
vations of the reserve (M. dobsoni, M. fotsifotsy,
M. gymnorhyncha, M. longicaudata, and M. sor-
icoides), and a few others occur from this zone
and up to the upper limit of forest cover on the
mountain (M. cowani, M. monticola, and M. prin-
cipula). M. gracilis is apparently restricted to the
upper montane zone of the Marojejy Massif.
However, this is based on a single capture of this
species in the reserve. The only species with an
apparently disjunct altitudinal distribution was
Oryzorictes hova, which was collected in lowland
forest at 450 m and in montane mossy forest at
1625 m. This latter result is almost certainly an
artifact of the aforementioned difficulty in trap-
ping Oryzorictes.

Among Microgale, there is no clear evidence
of species replacement along the elevational gra-
dient surveyed within the PN de Marojejy. At one
site inventoried over the past few years, the two
species of long-tailed shrew tenrecs, M. princi-
pula and the slightly smaller M. longicaudata, oc-
cur along the same slopes (Table 1 1-3). In the RS
d'Anjanaharibe-Sud, there is some evidence that
these two species might replace one another M.
principula was found only at 875 m and M. lon-
gicaudata at 1260-1950 m (Goodman & Jenkins,
1998). However, this pattern was not the case in
the PN de Marojejy; here, M. longicaudata was
recorded only at 1225 m, and M. principula oc-
curred higher on the mountain, at 1625-1875 m.
On the slopes of the PN d'Andohahela these two
species were broadly sympatric, with M. princi-
pula occurring at 440-1200 m and M. longicau-
data along the complete length of the transect, at
440-1875 m (Goodman et al., 1999a). Whether
these differences in the elevational distribution of
the two species are artifacts of the number of cap-

tures or reflect some aspect of habitat segregation
requires further study.

Comparison between the PN de Marojejy and
the RS d'Anjanaharibe-Sud Recent elevation
transects of Lipotyphla in several reserves in the
eastern humid forest biome, spanning the full lat-
itudinal breadth of the island from 12.5 to 24.5S,
provide an excellent means to study the relation-
ships between elevational and latitudinal gradi-
ents. If indeed there are latitudinal components
involved in these patterns, then before testing for
them it is important to look for consistency in
species composition and elevational range be-
tween sites at the same latitude. The PN de Ma-
rojejy and the RS d'Anjanaharibe-Sud provide the
opportunity for such a comparison. These two
sites are separated by approximately 40 km, and
the Andapa Basin lies between them. The summit
of Anjanaharibe Anivo (2064 m) is at 1444'S and
that of Marojejy (2033 m) is at approximately
1430'S. The botanical communities on these two
mountains show many similarities (Goodman &
Lewis, 1998; Chapter 3). A notable difference be-
tween them is that the summital zone of the Ma-
rojejy Massif contains a larger expanse of high-
elevation grassland than that of Anjanaharibe-
Sud. Furthermore, the Marojejy Massif lies closer
to the coast, which certainly has an effect on an-
nual rainfall (Donque, 1975); Marojejy is presum-
ably wetter.

During the survey of the PN de Marojejy, 13
species of shrew tenrecs and a mole tenrec belong-
ing to the subfamily Oryzorictinae were collected,
and two species of spiny tenrecs belonging to the
subfamily Tenrecinae were observed, giving a total
of 15 species of Tenrecidae. This is in comparison
with the RS d'Anjanaharibe-Sud, where 12 species
of Oryzorictinae and three species of Tenrecinae
(only one of which was collected) have been doc-
umented in this protected area (Goodman & Jen-
kins, 1998). Given that most of the Tenrecinae re-
cords for these two massifs are based on observa-
tions, we prefer to restrict our comparisons to ory-
zorictines. Of these animals, most species were
found in both the PN de Marojejy and the RS
d'Anjanaharibe-Sud (O. hova, M. cowani, M. dob-
soni, M. fotsifotsy, M. gymnorhyncha, M. longicau-
data, M. monticola, M. parvula, M. principula, M.
soricoides, and M. talazaci), but there are a few
anomalies, with M. brevicaudata and M. gracilis
collected only in PN de Marojejy and M. dryas
only in the RS d'Anjanaharibe-Sud. Microgale
brevicaudata seems to be limited to the coastal
plain and adjacent foothills (see pp. 208-209). For

222

FIELDIANA: ZOOLOGY

o

-o
-A

-IB---

Montagne d'Ambre

Andohahela

Marojejy

Andringitra

Anjanaharibe-Sud

CD
Q.
CO

Q.
CO

2

CD

2

o
5

u

9-

A

...-O

8 -
7-

O

6 -

a.

o- -

A .../'^^v ^N

\\

S,

4-

^

o s bb

3-

H

2 -

1 -

i

i l I I

i

250

250

500

750

1000 1250 1500

Elevation (m)

1750 2000

y-

8-

B

A:; ...

ft ' -"""

Q

6-

O'ffl

~ y/J^

3-

a./

2-

'""6

1 -

0-

i

ffl
I I I

i

i

500

750

1250 1500 1750 2000

Elevation (m)

Fig. 1 1-4. Plots of Lipotyphla species richness in relation to elevational transects for five different mountains on
Madagascar. Information is presented as all Tenrecidae species (A) and restricted to Microgale species (B). Data are
derived from the same sources mentioned in the footnotes to Table 11-3.

M. gracilis and M. dryas, the differences between
the sites is most likely an artifact of the difficulty
in trapping these species.

As far as elevational separation is concerned,
the altitudinal distribution of species is broadly
similar in both reserves, with M. parvula and M.
talazaci in lowland to montane zones and M. lon-
gicaudata, M. dobsoni, M. soricoides, M. gym-
norhyncha, M. cowani, and M. monticola in mon-
tane zones. M. principula is a notable exception,
since it was collected only in lowland areas in RS
d' Anjanaharibe-Sud and only in montane habitats
in PN de Marojejy. Thus, in general, the species
compositions and elevational distributions of li-
potyphlans, particularly Oryzorictinae, on these
two mountains are very similar.

Species Richness Over the past few years,
there has been a flurry of articles and hypotheses
to explain the distribution of plants and animals

along latitudinal and altitudinal gradients (e.g.,
Rahbek, 1997). Considerable attention has been
given to the effects of forest productivity in mold-
ing these patterns (Ricklefs & Schluter, 1993; Ro-
senzweig & Abramsky, 1993). A repeated pattern
of a species diversity gradient across a broad array
of taxa involves the peaking of species numbers
at middle elevations on mountains in tropical lat-
itudes (e.g., Janzen et al., 1976; Terborgh, 1977;
Heaney & Rickart, 1990; Olson, 1994; Rax worthy
et al., 1998); such a mid-elevational bulge pattern
may be related to measures of environmental
complexity and ecological productivity (see re-
view by Rosenzweig, 1992). In the PN
d' Andringitra, an analysis of soil invertebrates
showed that the elevational zone with the greatest
invertebrate density and taxonomic diversity was
in direct parallel with that portion of the mountain
with the highest Microgale species richness and

GOODMAN & JENKINS: LIPOTYPHLA

223

2000

1800-

1600-

E 1400-

.2 1200-

10 15 20 25

Latitude south

Latitude south

2000

1800-

1600-

1400-

5 1200-

1000

1000

Latitude south

1500 2000 2500

Summit of mountain (m)

3000

- Andohahela

- Marojejy

- Andringitra

- Anjanaharibe-Sud

- Montagne d'Ambre

Fig. 11-5. Plots assessing the relationships between various latitudinal and topographic variables and species
richness of Lipotyphla on Madagascar.

density (Goodman et al., 1996a). Thus, in this
case there is a correlation between productivity,
as measured by soil invertebrate density and di-
versity, and the distribution of Lipotyphla on
these slopes. We interpret this result as support
for the productivity hypothesis. Comparable data
are not available for the PN de Marojejy. but we
suspect the pattern found on the Andringitra Mas-
sif is applicable to Marojejy.

The distribution of Tenrecidae species in the
PN de Marojejy conforms to a unimodal. hump-
shaped pattern across the five altitudinal zones

surveyed (Fig. 1 1-4; Table 1 1-2). The highest spe-
cies richness of this family on the massif is nine
species, recorded at 1625 m, within the upper por-
tion of montane forest. When this measure is con-
fined to Microgale, both the 1225 and 1625 m
sites, each with eight species, constitute the zone
with the greatest diversity of shrew tenrecs. The
fewest species of Tenrecidae found in any zone
was four species at 775 m (transitional lowland-
montane forest) and 1875 m (montane/sclero-
phyllous forest). When this comparison is restrict-
ed to the genus Microgale, the pattern changes

224

FIELDIANA: ZOOLOGY

Table 1 1-5. Topographic settings in which Lipotyphla were captured by pitfall traps in the PN de Marojejy.

No. of captures at altitude and placement*

450 m

775 m

1225 m

1625 m

1875 m

Total

Species

VSR VSR

VSR VSR

Microgale brevicaudata
Microgale cowani
Microgale dobsoni
Microgale fotsifotsy
Microgale gymnorhyncha
Microgale longicaudata
Microgale monticola
Microgale parvula
Microgale principula
Microgale soricoides
Microgale talazaci
Oryzorictes hova
Total captures

3 1

112 4

16 3 4

3

1

3

5

2

4

4

6

2

1

9

9

9

1

1

5

1

1

1

6

1

1

2

2
1

1

1

2

2
1

1
1

5

4

2

4

2

3

6

10

4

1

2

1

2

1

1

1

3

6

3

4
3

4
1

1

2

4

2

1

2

3

3

6

2

1

2

1

2

2
2

3

11 19 15 12 9 15 5 6 1 33 40 36

* V = valley; S = slope; and R = ridge.

somewhat: the lowest species richness, at two, re-
mains the 775 m zone, followed by three species
in the lowland forest at 425 m, and finally by four
species at 1875 m. A parallel pattern of a mid-
elevational bulge in Tenrecidae species richness
has been found at several other mountains on the
island: the PN d'Andringitra (Goodman et al.,
1996a; Langrand & Goodman, 1997; Goodman,
unpubl. data), the RS d'Anjanaharibe-Sud (Good-
man & Jenkins, 1998), the PN d'Andohahela
(Goodman et al., 1999a), and the PN de la Mon-
tagne d'Ambre (Goodman et al., 1996a). The
sites, all inventoried by means of the same sam-
pling protocol, span the full latitudinal breadth of
the eastern humid forest biome in Madagascar,
from 12.5 to 24.5S. Further, the surveys of An-
dohahela, Andringitra, Anjanaharibe-Sud, and
Marojejy were conducted during the same period
(October to early December).

Although a mid-elevational bulge in species
richness recurs among these five mountains, there
is no consistent altitude at which diversity of Li-
potyphla peaks, whether examined across all of
the Tenrecidae (Fig. 11 -4 A) or restricted to Mi-
crogale spp. (Fig. 11-4B). Whether for the Ten-
recidae as a whole or specifically for the genus
Microgale, the zones that hold the largest number
of species vary from 1150 m on Montagne
d'Ambre to 1875 m on the Andohahela Massif.

There is no clear north-to-south trend sugges-
tive of elevational shifts in vegetational commu-
nities along a latitudinal gradient in the elevation
where maximum Tenrecidae species richness was
recorded (Fig. 1 1-5 A), in the total number of Ten-

recidae known from each mountain (Fig. 11-5B),
or in the total number of Microgale spp. recorded
on each mountain (Fig. 11 -5C). The elevation of
maximum diversity at Andohahela and Andringi-
tra, the two southern mountains closest to one an-
other in this sample, varies from 1210 m in An-
dringitra to 1875 m in Andohahela, and the inter-
mediate value between these two sites is from
Marojejy, toward the north end of the island. Fur-
ther, the Anjanaharibe-Sud and Marojejy Massifs,
within one degree of latitude of each other, show
divergent patterns in the zone of maximum spe-
cies richness.

Analyzing similar types of data to explain the
elevational distribution of Nesomyine rodents on
the slopes of the Montagne d'Ambre, Anjanahar-
ibe-Sud, Andringitra, and Andohahela Massifs,
Goodman et al. (1999b) found a very strong and
positive correlation between a mountain's abso-
lute height and the elevation with the most spe-
cies. However, for Tenrecidae, this pattern does
not hold (Fig. 11-5D). The strong correlation be-
tween these variables for Nesomyine rodents was
explained in that, on each of these mountains, the
zone showing the greatest species richness occurs
within the belt of wet montane rain forest. This
zone is wedged between the nearly perennial
cloud shroud in the summital zone, with low lev-
els of direct sunlight and with problems associated
with waterlogging, and the lowland area, which
has greater direct solar radiation, although, at least
seasonally, it experiences higher evaporation rates
and desiccation associated with water shortage.
We strongly suspect that this same pattern holds

GOODMAN & JENKINS: LIPOTYPHLA

225

for the Tenrecidae, but we lack the statistical
backing that was found for the rodents.

Habitat Preferences Pitfall traps were
placed in three different microhabitats at each el-
evation: in valley bottoms, on the slopes of hills,
and on ridge crests. These three topographical
zones generally have distinctly different botanical
communities (see Chapter 3). The distribution of
oryzorictines was roughly equal in all three hab-
itats, with 33 individuals trapped in valley bot-
toms, 40 on slopes, and 36 on ridges (Table 1 1 -
5). This is in contrast to the other sites, where
Microgale were captured more frequently in val-
leys than on slopes or ridges (Goodman et al.,
1996a).

At the species level, there is little evidence of
differential captures in any of these three micro-
habitats, but only one species, M. monticola, was
captured in sufficient numbers to properly assess
such preferences. Approximately 50% more in-
dividuals of this species were captured in the PN
de Marojejy on slopes as compared with valleys
and ridges. In the nearby RS d'Anjanaharibe-Sud,
the same basic pattern exists: about half the num-
ber of this species were captured on ridges and in
valleys as compared with slopes (Goodman &
Jenkins, 1998).

Acknowledgments

This field work could not have been accom-
plished without the help of the staff of the World
Wide Fund for Nature, Andapa, particularly Jean
Marc Garreau and Julien Befourouack. We are
grateful to officials of the DEF and ANGAP for
permits to enter the reserve and collect specimens.
Laurent Granjon of the Museum National
d'Histoire Naturelle, Paris, and Daniel Rakoton-
dravony of the Departement de Biologie Animale,
Universite d' Antananarivo, kindly made speci-
mens available for this study. For comments on
an earlier version of this chapter we are grateful
to Link Olson, Howard Whidden, and an anony-
mous reviewer.

Literature Cited

Ade, M. 1996. Morphological observations on a Micro-
gale specimen (Insectivora, Tenrecidae) from western
Madagascar, pp. 251-255. In Ganzhorn, J. U., and J.-
P. Sorg, eds., Ecology and Economy of a Tropical Dry

Forest in Madagascar. Primate Report, special issue,
Deutsches Primatenzentrum, Gottingen.

Albignac, R. 1970. Mammiferes et oiseaux du massif
du Tsaratanana (Madagascar Nord). Memoires OR-
STOM, 37: 223-229.

Butler, P. M., and M. Greenwood. 1979. Soricidae
(Mammalia) from the Olduvai Gorge, Tanzania. Zoo-
logical Journal of the Linnean Society, 67: 329-379.

Carleton, M. D., and D. F. Schmidt. 1990. Systematic
studies of Madagascar's endemic rodents (Muroidea:
Nesomyinae): An annotated gazetteer of collecting lo-
calities of known forms. American Museum Novita-
tes, 2987: 1-36.

Dewar, R., and S. Rakotovololona. 1992. La chasse
aux subfossiles: Les preuves du onzieme siecle au trei-
zieme siecle. Taloha, 11: 4-15.

Donque, G. 1975. Contribution geographique e l'etude
du climat de Madagascar. Nouvelle Imprimerie des
Arts Graphiques, Antananarivo, 478 pp.

Duckworth, J. W. 1990. Systematic list of mammals
recorded in Marojejy, pp. 1 14-129. In Safford, R., and
W. Duckworth, eds., A Wildlife Survey of Marojejy
Nature Reserve, Madagascar. Study Report no. 40. In-
ternational Council for Bird Preservation, Cambridge,
171 pp.

Eisenberg, J. E, and E. Gould. 1970. The tenrecs: A
study in mammalian behavior and evolution. Smith-
sonian Contributions to Zoology, 27: 1-137.

Ganzhorn, J. U., S. Sommer, J. -P. Abraham, M. Ade,
B. M. Raharivololona, E. R. Rakotovao, C. Rak-
otondrasoa, and R. Randriamarosoa. 1996. Mam-
mals of the Kirindy Forest with special emphasis on
Hypogeomys antimena and the effects of logging on
the small mammal fauna, pp. 215-232. In Ganzhorn,
J. U., and J. -P. Sorg, eds., Ecology and Economy of
a Tropical Dry Forest in Madagascar. Primate Report,
special issue, Deutsches Primatenzentrum, Gottingen,
46-1: 1-382.

Goodman, S. M., A. Andrianarimisa, L. E. Olson, and
V. Soarimalala. 1996b. Patterns of elevational dis-
tribution of birds and small mammals in the humid
forests of Montagne d'Ambre. Madagascar. Ecotropi-
ca, 2: 87-98.

Goodman, S. M., M. D. Carleton, and M. Pidgeon.
1 999b. The rodents of the Reserve Naturelle Integrale
d'Andohahela, Madagascar, pp. 217-249. In Good-
man, S. M., ed., A Floral and Faunal Inventory of the
Reserve Naturelle Integrale d'Andohahela, Madagas-
car: With Reference to Elevational Variation. Fieldi-
ana: Zoology, n.s., 94: 1-297.

Goodman, S. M., J.-M. Duplantier, P. J. Rakotomal-
aza, A. P. Raselimanana, R. Rasoloarison, M. Ra-
vokatra, V. Soarimalala, and L. Wilme. 1998b. In-
ventaire biologique de la foret d'Ankazomivady, Am-
bositra. Akon'ny Ala, 24: 19-32.

Goodman, S. M., J. U. Ganzhorn, L. E. Olson, M.
Pidgeon, and V. Soarimalala. 1997. Annual varia-
tion in species diversity and relative density of rodents
and insectivores in the Pare National de la Montagne
d'Ambre, Madagascar. Ecotropica, 3: 109-118.

Goodman, S. M., and P. D. Jenkins. 1998. The insec-
tivores of the Reserve Speciale d'Anjanaharibe-Sud,
pp. 139-161. In Goodman, S. M., ed., A Floral

226

FIELDIANA: ZOOLOGY

and Faunal Inventory of the Reserve Speciale
d'Anjanaharibe-Sud, Madagascar: With Reference to
Elevational Variation. Fieldiana: Zoology, n.s., 90: 1-
246.

Goodman, S. M., R D. Jenkins, and M. Pidgeon. 1999a.
The Lipotyphla (Tenrecidae and Soricidae) of the Re-
serve Naturelle Integrale d'Andohahela, Madagascar,
pp. 187-216. In Goodman, S. M., ed., A Floral and
Faunal Inventory of the Reserve Naturelle Integrale
d'Andohahela, Madagascar: With Reference to Ele-
vational Variation. Fieldiana: Zoology, n.s., 94: 1-
297.

Goodman, S. M., and B. A. Lewis. 1998. Description
of the Reserve Speciale d'Anjanaharibe-Sud, Mada-
gascar, pp. 9-16. In Goodman, S. M., ed., A Floral
and Faunal Inventory of the Reserve Speciale
d'Anjanaharibe-Sud, Madagascar: With Reference to
Elevational Variation. Fieldiana: Zoology, n.s., 90: 1-
246.

Goodman, S. M., D. Rakotondravony, L. E. Olson, E.
Razafimahatratra, and V Soarimalala. 1998a. Les
insectivores et les rongeurs, pp. 80-93. In Rakoton-
dravony, D., and S. M. Goodman, eds., Inventaire
Biologique Foret d'Andranomay, Anjozorobe.
World Wide Fund for Nature-Centre d'lnformation et
de Documentation Scientifique et Technique, Anta-
nanarivo, Recherches pour le Developpement. Serie
Sciences Biologiques, no. 13: 1-110.

Goodman, S. M., D. Rakotondravony, G. Schatz, and
L. Wilme. 1996c. Species richness of forest-dwelling
birds, rodents and insectivores in a planted forest of
native trees: a test case from the Ankaratra, Madagas-
car. Ecotropica, 2: 109-120.

Goodman, S. M., C. J. Raxworthy, and R D. Jenkins.
1996a. Insectivore ecology in the Reserve Naturelle
Integrale d'Andringitra, Madagascar, pp. 218-230. In
Goodman, S. M., ed., A Floral and Faunal Inventory
of the Eastern Slopes of the Reserve Naturelle Inte-
grale d'Andringitra, Madagascar: With Reference to
Elevational Variation. Fieldiana: Zoology, n.s., 85: 1-
319.

Grandidier, A. 1870. Description de quelques animaux
nouveaux, decouverts a Madagascar, en novembre
1 869. Revue et Magasin de Zoologie, serie 2, 22: 49-
50.

Grandidier, G. 1899. Description d'une nouvelle espece
d' insectivore provenant de Madagascar. Bulletin du
Museum d'Histoire Naturelle 5: 349.

. 1934. Deux nouveaux mammiferes insectivores

de Madagascar, Microgale drouhardi et M. parvula.
Bulletin du Museum National d'Histoire Naturelle,
serie 2, 6: 474-477.

Grandidier, G., and G. Petit. 1930. Description d'une
espece nouvelle d'lnsectivore Malgache, suivie de re-
marques critiques sur le genre Oryzoryctes. Bulletin
du Museum National d'Histoire Naturelle, serie 2, 2:
498-505.

. 1931. Un type nouveaux de centetide malgache,

Paramicrogale occidentalis. Bulletin de la Societe
Zoologique de France, 56: 126-139.

Griveaud, P. 1960. line mission entomologique au Ma-
rojejy. Naturaliste Malgache, 12: 43-55.

Heaney, L. R., and E. A. Rickart. 1990. Correlations

of clades and clines: Geographic, elevational, and
phylogenetic distribution patterns among Philippine
mammals, pp. 321-332. In Peters, G., and R. Hutterer,
eds., Vertebrates in the Tropics. Museum Alexander
Koenig, Bonn.

Humbert, H. 1951. Les territoires phytogeographiques
du nord de Madagascar. Compte Rendus des Seances
de la Societe de Biogeographie, 246: 176-184.

Janzen, D. H., D. M. Ataroff, M. Farinas, S. Reves,
N. Rincon, A. Soler, P. Soriano, and M. Vera. 1976.
Changes in the arthropod community along an ele-
vational transect in the Venezuelan Andes. Biotropica,
8: 193-203.

Jenkins, P. D. 1988. A new species of Microgale (In-
sectivora: Tenrecidae) from northeastern Madagascar.
American Museum Novitates, 2910: 1-7.

. 1993. A new species of Microgale (Insectivora:

Tenrecidae) from eastern Madagascar with an unusual

dentition. American Museum Novitates, 3067: 1-11.

Jenkins, P. D., S. M. Goodman, and C. J. Raxworthy.

1996. The shrew tenrecs {Microgale) (Insectivora:
Tenrecidae) of the Reserve Naturelle Integrale
d'Andringitra, Madagascar, pp. 191-217. In S. M.
Goodman, ed., A Floral and Faunal Inventory of the
Eastern Slopes of the Reserve Naturelle Integrale
d'Andringitra, Madagascar: With Reference to Ele-
vational Variation. Fieldiana: Zoology, n.s., 85: 1-
319.

Jenkins, P. D., C. J. Raxworthy, and R. A. Nussbaum.

1997. A new species of Microgale (Insectivora, Ten-
recidae), with comments on the status of four other
taxa of shrew tenrecs. Bulletin of the Natural History
Museum London (Zoology), 63: 1-12.

Kaudern, W 1918. Quartare Fossilien aus Madagascar.
Zoologische Jahrbiicher, Abteilung fur Systematik,
Geographie und Biologie der Tiere, 41: 519-533.

Langrand, O., and S. M. Goodman. 1997. Inventaire
biologique des oiseaux et des micro-mammiferes des
zones sommitales de la Reserve Naturelle Integrale
d'Andringitra. Akon'ny Ala 20: 39-54.

MacPhee, R. D. E. 1986. Environment, extinction, and
Holocene vertebrate localities in southern Madagascar.
National Geographic Research, 2: 441-455.

. 1987. The shrew tenrecs of Madagascar: Sys-
tematic revision and Holocene distribution of Micro-
gale (Tenrecidae, Insectivora). American Museum
Novitates, 2889: 1-45.

Major, C. I. Forsyth. 1896. Diagnoses of new mam-
mals from Madagascar. Annals and Magazine of Nat-
ural History, series 6, 18: 318-325.

. 1897. On the general results of a zoological

expedition to Madagascar in 1894-96. Proceedings of
the Zoological Society of London (for 1896): 971
981.

Mills, J. R. E. 1966. The functional occlusion of the
teeth of Insectivora. Journal of the Linnean Society
(Zoology), 47: 1-125.

Nicoll, M. E., and O. Langrand. 1989. Madagascar:
Revue de la conservation et des aires protegees. World
Wide Fund for Nature, Gland, Switzerland, 374 pp.

Nicoll, M. E., and G. B. Rathbun. 1990. African In-
sectivora and Elephant-Shrews: An action plan for
their conservation. International Union for the Con-

GOODMAN & JENKINS: LIPOTYPHLA

227

servation of Nature and Natural Resources, Gland,
Switzerland, 53 pp.

Olson, D. M. 1994. The distribution of leaf litter inver-
tebrates along a Neotropical altitudinal gradient. Jour-
nal of Tropical Ecology, 10: 129-150.

Rahbek, C. 1997. The relationship among area, eleva-
tion, and regional species richness in Neotropical
birds. The American Naturalist, 149: 875-902.

Randrianjafy Rasoloarisoa, V. N. R. 1993. Contri-
bution a l'etude bio-ecologique du peuplement de mi-
cromammiferes dans la foret de l'Ankarafantsika. Me-
moire de Diplome d' Etudes Approfondies, Sciences
Biologiques Appliquees. Ecologie Environnementale,
Universite d' Antananarivo, Antananarivo.

Raxworthy, C. J., and R. A. Nussbaum. 1994. A rain-
forest survey of amphibians, reptiles and small mam-
mals at Montagne d'Ambre, Madagascar. Biological
Conservation, 69: 65-73.

Raxworthy, C. J., F. Andreone, R. A. Nussbaum, N.
Rabibisoa, and H. Randriamahazo. 1998. Amphibi-
ans and reptiles of the Anjanaharibe-Sud Massif: Ele-
vational distribution and regional endemicity, pp. 79-
92. In Goodman. S. M., ed.. A Floral and Faunal In-
ventory of the Reserve Speciale d'Anjanaharibe-Sud.
Madagascar: With Reference to Elevational Variation.
Fieldiana: Zoology, n.s., 90: 1-246.

Ricklefs. R. E., and D. Schluter, eds. 1993. Species
diversity in ecological communities. University of
Chicago Press, Chicago. 414 pp.

Rosenzweig, M. L. 1992. Species diversity gradients:
We know more and less than we thought. Journal of
Mammalogy, 73: 715-730.

Rosenweig, M. L., and Z. Abramsky. 1993. How are
diversity and productivity related?, pp. 52-65. In
Ricklefs, R. E., and D. Schluter, eds., Species Diver-
sity in Ecological Communities. University of Chi-
cago Press. Chicago, 414 pp.

Stephenson, P. J. 1994. Seasonality effects on small
mammal trap success in Madagascar. Journal of Trop-
ical Ecology, 10: 439-444.

. 1995. Small mammal microhabitat use in low-
land rain forest of northeast Madagascar. Acta Ther-
iologica. 40: 425-438.

Swindler, D. R. 1976. Dentition of Living Primates.
Academic Press, London, 308 pp.

Terborgh, J. 1977. Bird species diversity along an An-
dean elevational gradient. Ecology, 58: 1007-1019.

Thomas. (M. R.) Oldfield. 1882. Description of a new
genus and two new species of Insectivora from Mad-
agascar. Journal of the Linnean Society (Zoology). 16:
319-322.

. 1884. Description of a new species of Micro-
gale. Annals and Magazine of Natural History, series
5. 14: 337-338.

. 1918. On the arrangement of the small Tenre-

cidae hitherto referred to Oryzorictes and Microgale.
Annals and Magazine of Natural History, series 9, 1:
302-307.

. 1926. On some small mammals from Madagas-
car. Annals and Magazine of Natural History, series
9. 17: 250-252.

Viette. P. 1991. Principales localities ou des Insectes ont

ete recueillis a Madagascar. Faune de Madagascar,
supplement 2. Private Printing.
Voss, R. S., and L. H. Emmons. 1996. Mammalian di-
versity in Neotropical lowland rainforests: A prelim-
inary assessment. Bulletin of the American Museum
of Natural History, 230: 1-115.

Key 1. The genera of Tenrecidae occurring in
PN de Marojejy.

1. Pelage spinous; tail very short 2

Pelage soft, lacking spines; tail medium to long

4

2. Close-set, sharp spines cover dorsal surface

Setifer

Dorsal surface covered with a mixture of
spines and long, coarse hair 3

3. Dorsal pelage dark with longitudinal pale
stripes; head and body length < 200 mm . . .

Hemicentetes*

Dorsal pelage uniform brown; head and body
length > 260 mm Tenrec

4. Body robust, forelimbs robust, forefeet broad
with enlarged, stout claws; C longer than II

Oryzorictes

Body slender to moderately robust, forelimbs
not enlarged, forefeet slender to moderately
broad, claws short to moderately lengthened;
II longer or subequal in length to C

Microgale

* Hemicentetes was neither collected nor
observed during the survey, but it has been
recorded previously from this reserve. For fur-
ther comments, see Discussion (p. 221).

Key 2. The species of Microgale occurring in
PN de Marojejy.

1. Size very small, HB < 62, CIL < 17.1

M. parvula

Size larger, HB > 65, CIL > 20.0 2

2. Ratio of TL:HB > 2.0 3

Ratio of TL:HB < 1.5 4

3. Size smaller. HB < 70, CIL < 21.0

M. longicaudata

Size larger, HB > 70, CIL > 21.0

M. principula

4. Digits and tail tip contrastingly paler than
body, tail, and feet M. fotsifotsy

228

FIELDIANA: ZOOLOGY

Tail tip and digits not obviously paler than
rest of body 5

5. Proboscis long, large rhinarium extends pos-
terodorsally onto muzzle; forefeet broad, fo-

reclaws enlarged 6

Small rhinarium confined to anterior of short
proboscis; forefeet slender without length-
ened foreclaws 7

6. Posterior region of rhinarium with transverse
striae; ratio of I-P3 : UTL < 0.55

M. gymnorhyncha

Posterior region of rhinarium reticulated; ra-
tio of I-P3 : UTL > 0.57 M. gracilis

7. Size smaller: CIL < 24; tail shorter < 75 . . 8
Size larger: CIL > 24; tail longer > 88 . . 9

8. Tail shorter: ratio of TL : HB < 0.6

M. brevicaudata

Tail longer: ratio of TL : HB > 0.8

M. cowani

9. Size smaller: HB < 100, CIL < 26.0 . . 10

Size larger: HB > 100, CIL > 30.0 .... 11

10. Pelage dark brown dorsally, slightly lighter
ventrally; TL longer than HB, ratio > 1.2 . .

M. monticola

Pelage lightish buff brown dorsally, reddish
buff ventrally; TL subequal to HB, ratio <
1.1 M. soricoides

11. Cranial size smaller, CIL < 32.0; body size
averaging smaller, HB < 108, WT < 32 ...

M. dobsoni

Cranial size larger, CIL > 34.0; body size av-
eraging larger, HB > 104, WT > 33

M. talazaci

GOODMAN & JENKINS: LIPOTYPHLA

229

Chapter 12

Rodents of the Pare National de
Marojejy, Madagascar

Michael D. Carleton 1 and Steven M. Goodman 2

Abstract

The small mammal inventory of the Pare National de Marojejy, conducted between 4 October
and 20 November, 1996, in five elevational stations from 450 to 1875 m, produced vouchered
evidence of seven species of native rodents (Muridae: Nesomyinae) that inhabit the park
Eliurus grandidieri, E. majori, E. minor, E. tanala, E. xvebbi, Gymnuromys roberti, and Voalavo
gymnocaudus. Regular sightings of a large Nesomys (probably N. rufus) suggest the presence
of this diurnal form, yet its numbers were low and none were live trapped. The introduced
rodent Rattus rattus (Muridae: Murinae) was also collected, but unlike in field surveys of
several other northern forest sites, it proved to be uncommon.

The variable incidence of white tail tips in the series of E. majori from Marojejy prompted
taxonomic review of the status of E. penicillatus, a nominal species known only from its type
locality in east-central Madagascar and uniformly possessing a white caudal tip. Morphometric
evaluation, using craniodental data, of E. penicillatus and six E. majori samples across much
of its known range recommends that the two should be maintained as separate species. Com-
parable analyses of nine population samples of E. webbi from throughout its range uncovered
only minor differences consistent with geographic variation within a single, widely distributed
species.

The highest diversity of native species was documented at 1250 m, near the lower limit of
montane forest (six species); fewer species occupied lowland rain forest, at 450 m (one), and
sclerophyllous montane forest, at 1875 m (three). The sympatric coincidence, elevational limits,
and ecological agreement of native species on the massifs of Marojejy and nearby Anjanahar-
ibe-Sud are strongly concordant and reaffirm broad distributional themes of nesomyines within
the eastern humid forest biome. Although certain nesomyine species do occur in sclerophyllous
montane forest up to tree line, none has been recovered from the ericaceous grassland and bush
near the summits of Marojejy and Anjanaharibe-Sud. The apparent absence of native rodents
in comparable alpine habitat on these northern highlands and the presence of one genus (Brach-
yuromys) in similar settings on the Central High Plateau are discussed.

Resume

L'inventaire des petits mammiferes du Pare National de Marojejy, conduit entre le 4 octobre
et le 20 novembre 1996, dans cinq stations reparties a des altitudes de 450 a 1875 metres,
permit d'obtenir des donnees de reference sur sept especes des rongeurs indigenes (Muridae:
Nesomyinae) distributes dans le pare Eliurus grandidieri, E. majori, E. minor, E. tanala, E.

1 Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560,
U.S.A.

2 Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605-2496, U.S.A.

CARLETON & GOODMAN: RODENTS 231

webbi, Gymnuromys roberti et Voalavo gymnocaudus. Des observations regulieres d'un grand
Nesomys (probablement N. rufus) confirment la presence de cette forme diurne bien que ses
effectifs etaient bas et qu'aucun individu n'a ete capture vivant. Le rongeur introduit Rattus
rattus (Muridae: Murinae) a egalement ete capture mais contrairement aux inventaires de terrain
menes dans d'autres sites forestiers du nord, il s'est ici avere rare.

L' incidence variable du blanc sur le bout de la queue de la serie des E. majori obtenue du
Marojejy incita a la revision taxinomique du statut de l'espece nominative E. penicillatus,
uniquement connue de la localite du type dans le centre est de Madagascar, et qui montre
invariablement une queue terminee de blanc. L'evaluation morphometrique, en utilisant des
donnees craniodentales de E. penicillatus et de six echantillons d'. majori dont les provenance
couvrent la plupart de son aire connue de distribution, montrent que les deux taxons devraient
etre considered comme deux especes distinctes. Des analyses comparables effectuees sur neuf
echantillons d'. webbi provenant de localites couvrant son aire de distribution ne montrerent
que des differences mineures en rapport avec une variation geographique au sein d'une espece
a large distribution.

La diversite d'especes indigenes la plus importante a ete relevee a 1250 m d'altitude pres
de la limite inferieure de la foret de montagne (six especes), peu d'especes etant rencontrees
dans la foret humide de basse altitude a 450 m (une) ou dans la foret sclerophylle de montagne
a 1875 m (trois). La coincidence dans la sympatrie, les limites altitudinales. et les associations
ecologiques des especes indigenes des massifs du Marojejy et d'Anjanaharibe-Sud voisin con-
cordent nettement et reaffirment les grands themes de la distribution des Nesomyinae dans le
biome de la foret humide de Test. Si certaines especes de Nesomyinae sont rencontrees dans
la foret sclerophylle de montagne jusqu'a la lisiere superieure, aucune espece n'a ete capturee
dans les zones sommitales ericoides, couvertes d'herbes et de buissons, des massifs du Marojejy
et d'Anjanaharibe-Sud. L'apparente absence de rongeurs indigenes dans l'habitat propice des
sommets de ces hautes montagnes du nord et la presence d'un genre (Brachyuromys) dans des
milieux similaires du Haut Plateau Central sont discutees.

Introduction

Between 1950 and 1985, approximately 35% of
Madagascar's eastern humid forest was cleared
(Green & Sussman, 1990), a trend that continues
at an alarmingly high rate. The eastern humid for-
est (sensu Humbert, 1965) formerly covered much
of Madagascar's eastern versant, forming a nearly
unbroken swath of lowland and montane tropical
vegetation that stretched nearly 1,500 km in
length and extended inland from the coast, up the
eastern escarpment, and onto the central high-
lands, a vertical step of at least 1 ,000 m. The ma-
jority of primary forest in eastern Madagascar
now occurs only as disjunct parcels, often situated
within the boundaries of protected areas. Infor-
mation on the biota of these isolates is therefore
important to approximate the original fauna of the
eastern humid forest biome and to understand dis-
tributional patterns within it.

This report focuses on the endemic rodents
(Muridae: Nesomyinae) of the Pare National (PN)
de Marojejy (14.5 south latitude) in the northern
highlands, as revealed by a field investigation un-

dertaken between 4 October and 20 November
1996 by Goodman. This contribution forms an-
other part of the renewed inventory of small
mammals occurring along the forested slopes of
several mountains in eastern Madagascar: PN de
la Montagne d'Ambre, 12.5 south latitude
(Goodman et al., 1996, 1997), Reserve Speciale
(RS) d'Anjanaharibe-Sud, 14S (Carleton &
Goodman, 1998; Goodman & Carleton, 1998), the
Reserve Naturelle Integrate (RNI) d'Andringitra,*
22S (Goodman & Carleton, 1996; Goodman &
Rasolonandrasana, in press), and the RNI
d'Andohahela, 25S (Goodman, Carleton, & Pidg-
eon, 1999). The same kinds of traps and sampling
regimen were employed at each of these sites, and
the surveys were, for the most part, conducted
during the same season. Collectively, the five sites
span the complete latitudinal breadth of the is-
land's eastern forests. Special attention is devoted
to comparison of the rodent faunas of the PN de

* Since the publication of these faunal reports the sta-
tus of the Andringitra and Andohahela reserves has been
changed to Pare National.

232

FIELDIANA: ZOOLOGY

Marojejy and the RS d'Anjanaharibe-Sud, which
represent the reserves with the most complete fau-
nal documentation in the northern highlands.

Previous Work in the Region

The Marojejy Massif has been the subject of
numerous zoological inventories, most of them
entomological (Griveaud, 1960; Guillaumet et al.,
1975), but little information based on preserved
and critically identified vouchers is available on
the rodents occurring in the park.

Between 1929 and 1931, the Mission Zoolo-
gique Franco-Anglo-Americaine (MZFAA) con-
ducted an extensive inventory of the birds of
Madagascar and incidentally collected mammal
specimens. Their closest approach to the Marojejy
Massif was a highland site, 1800 m, about "one
day west of Andapa" (Rand, 1936), which Jen-
kins (1987) reckoned as 1439'S, 4922'E. Al-
though a single elevation was officially recorded,
the birds collected appear to represent a mixture
of highland and lowland forms. To judge from
elevational ranges of the avifauna from other
well-documented mountains, however, it appears
that the MZFAA collectors gathered specimens
from a broad range of habitats and altitudes (Haw-
kins et al., 1998). The few small mammals ob-
tained near the Andapa site are Rattus [rattus],
Eliurus minor, and Nesomys rufus (Rand, 1932;
Carleton & Schmidt, 1990), and these species are
generally known to have broad elevational ranges
in the region (Goodman & Carleton, 1998), in-
cluding highland areas. Thus, the elevational oc-
currence of mammals reported from the site sug-
gests no distributional irregularities such as are
suspected for the bird data. The possibility that
MZFAA material may have been collected over a
wide elevational range is pertinent to interpreta-
tion of altitudinal records based on specimens ob-
tained by their expedition.

In November 1958, an entomological mission
to the Marojejy Massif was organized by the In-
stitut de Recherche Scientifique de Madagascar
(IRSM), under the direction of R. Paulian and
with field supervision by R Griveaud. The general
results of the mission presented a list of small
mammals observed but did not mention any ro-
dent species (Griveaud, 1960).

In 1988, a student research group from Cam-
bridge University, working with Malagasy coun-
terparts, conducted small mammal trapping with

Sherman traps in several areas on the Marojejy
Massif (Duckworth & Rakotondraparany, 1990).
In the southeastern portion of the PN de Marojejy
(then still classified as an RNI), the same area
where Goodman worked in 1996, the team cap-
tured two Eliurus cf. myoxinus at 300 m (170
trap-nights) and six more individuals at 1300 m
(165 trap-nights). Unfortunately, no voucher spec-
imens were preserved, and the species taxonomy
of the genus has changed dramatically in the past
few years (Carleton, 1994). No specific determi-
nations can be reliably made on the basis of the
external measurements provided (Duckworth &
Rakotondraparany, 1990, p. 128). This group also
reported sight observations for Brachytarsomys
albicauda in the southeastern portion of the re-
serve (700 m) and for Nesomys in both the south-
western (800 m) and northwestern sectors (be-
tween 1000 and 1400 m). In their summary of
mammals known from the reserve, Nicoll and
Langrand (1989) listed the rodent species as re-
ported by the Cambridge University expedition.

In 1994, a multidisciplinary biological inven-
tory was conducted in the RS d'Anjanaharibe-
Sud, just to the west of the Marojejy Massif and
Andapa Basin. The rodent fauna was documented
in a monograph that included the description of a
new genus and species, Voalavo gymnocaudus, as
well as a new species of Eliurus, E. grandidieri
(Carleton & Goodman, 1998), and a general re-
view of the reserve's nine species of native ro-
dents (Goodman & Carleton, 1998). Since the
field techniques used during the 1996 inventory
of the PN de Marojejy were identical to those em-
ployed in the nearby RS d'Anjanaharibe-Sud, we
make regular comparisons between these two re-
serves throughout the chapter.

Materials and Methods

This study is based on fieldwork conducted be-
tween 4 October and 20 November 1996 by
Goodman. Carleton verified taxonomic determi-
nations and undertook systematic comparisons.

Field Methods and Trapping Protocol

The general field protocols followed those pre-
viously outlined in parallel reports on the rodents
of the RNI d'Andringitra (Goodman & Carleton,
1996), the RS d'Anjanaharibe-Sud (Goodman &

CARLETON & GOODMAN: RODENTS

233

Carleton, 1998), and the RNI d'Andohahela
(Goodman, Carleton, & Pidgeon, 1999). In the PN
de Marojejy, five altitudinal zones were sampled
(450, 775, 1250, 1625, and 1875 m), representing
the gamut of principal vegetational formations
known in the region, from lowland forest through
sclerophyllous forest to open savanna above tree
line (see Chapters 1 and 3). The area surveyed at
450 m clearly showed signs of recent human dis-
turbance, those at 775 and 1250 m much less so,
while the two uppermost sites seemed pristine.

At each of the five elevations, trap lines were
maintained for a minimum of six nights (Table 1 2-
1). Each trap line, numbered sequentially starting
with the 450 m zone, consisted of Sherman live
traps (9 X 3.5 X 3 inch) and National live traps
(16 X 5 X 5 in.) in a ratio of 4:1. Traps were
baited daily, generally between 1500 and 1700
hours, with finely ground peanut butter. They
were inspected at least twice per day, once at
dawn and again in late afternoon. At each site,
sampling was also conducted with pitfall traps,
but this technique yielded few rodents (see Chap-
ter 11), and these incidental captures were not in-
cluded in calculations of trapping success and bio-
mass.

A trap-night is defined as one live trap in use
for a 24-hour period (dawn to dawn). The total
number of trap-nights accrued in each elevation
varied slightly; consequently, the first 500 trap-
nights in an elevational zone are considered the
"standardized" trapping regimen in order to fa-
cilitate comparisons among the sites sampled.
Standing biomass of a species is based on the total
catch of individuals during a standardized trap-
ping regimen multiplied by average body weight
of the species (see Table 12-5). We depended ex-
clusively on live trap techniques during this in-
ventory for reasons explained previously by
Goodman and Carleton (1998, p. 201).

To quantify differences in spatial distribution of
small mammal captures, several trapping vari-
ables were systematically recorded for each trap
installed: (1) type of trap, (2) total length of trap
line, (3) distance between traps, and (4) specific
placement of trap, including its substrate, sur-
rounding forest structure, and position on or
height above the ground. A system for the cate-
gorization of microhabitat used during this survey
is identical to that presented by Goodman and
Carleton (1998), as follows:

On Ground ( 1 ) in leaf litter, generally in area
of open understory; (2) under decomposed
downed trees or woody vegetation; (3) by tree

root or trunk, with or without cavity or hole; (4)
miscellaneous, including placement under ex-
posed rocks or boulders, at base of rock face, at
entrance of hole in ground, in thick herbaceous
vegetation or dense shrubbery, or on moss-cov-
ered rocks.

Above Ground (1') on liana, limb, or trunk
<10 cm diameter in horizontal to vertical posi-
tion; (2') on liana, limb or trunk >10 cm diameter
in horizontal to vertical position; (3') on limbs or
trunks suspended by lianas; (4') miscellaneous,
including placement on bamboo stalks, in small
cavities at junctions of tree limbs, or on large
moss-covered rocks.

Specimens and Measurements

Captured animals were prepared either as stan-
dard museum skins with associated skulls and
partial skeletons, or as fluid-preserved carcasses
(some with skulls removed), or as full skeletons.
Specimens prepared as whole carcasses were
wrapped in fine cheesecloth before immersion in
formalin to prevent loss or mixing of ectoparasites
between their specific hosts (see Chapter 7). Near-
ly all rodents captured during our work in the PN
de Marojejy were prepared as vouchers. This ma-
terial is housed in the Field Museum of Natural
History (FMNH) and Departement de Biologie
Animale, Universite d' Antananarivo (UADBA).
Specimens deposited immediately after the survey
in the latter institution have not yet been cata-
logued and are individually referenced by the col-
lector's field numbers (UA-SMG). To confirm tax-
onomic identifications, nesomyine holdings in
other museums (see Appendix in Goodman &
Carleton, 1996) were also consulted.

Six measurements, in millimeters (mm) or
grams (g), were taken by Goodman for each spec-
imen in the flesh. Measurement abbreviations and
definitions are given below.

TOTL (total length of body and tail): from the
tip of the nose to the end of the last cau-
dal vertebra (not including terminal hair
tuft)

HBL (head and body length): from the tip of
the nose to the distalmost point of the
body (at base of tail)

TL (tail length): from the base of the tail

(held at right angle to the body) to the
end of the last caudal vertebra (not in-
cluding terminal hair tuft)

234

FIELDIANA: ZOOLOGY

Table 12-1. Summary of five trap lines (each with 100 live traps) in the PN de Marojejy.*

Elevation

Length (m)
of line

Mean distance (m)
between traps

No. of traps
aboveground

Mean height (m)
aboveground

450 m (5-12 Oct)
Line 1

795

7.9 4.96 (1-21)

31

1.3 0.58 (0.2-2.5)

775 m (15-23 Oct)
Line 2

575

5.8 3.20 (2-16)

29

1.6 0.28 (0.2-2.5)

1250 m (26 Oct-1 Nov)
Line 3

575

5.9 4.26 (1-26)

33

1.7 0.71 (0.5-3)

1625 m (6-12 Nov)
Line 4

385

3.9 2.34 (1-9)

28

1.7 0.65 (0.2-3)

1875 m (13-19 Nov)
Line 5

540

5.6 3.50 (1-20)

8

1.3 0.62(0.2-2)

* Each line consisted of National and Sherman live traps in a ratio of 4: 1 (see p. 234).
Descriptive statistics are presented as mean SD (and range).

HFL (hind foot length): from the heel to the
tip of the longest toe (not including claw)

EL (ear length): from the basal notch to the

distal tip of the pinna

WT (weight): measured with Pesola spring
scales, to 0.5 g for animals <100 g
and to 1.0 g for those between 101 and
300 g.

Sixteen cranial and two dental dimensions were
measured by Carleton to the nearest 0. 1 mm using
handheld digital calipers accurate to 0.03 mm.
These measurements, and their abbreviations, fol-
low the anatomical landmarks defined and illus-
trated in Carleton (1994).

BBC, breadth of the braincase

BIF, breadth of incisive foramina

BMls, breadth of the bony palate across the first

upper molars
BOC, breadth across the occipital condyles
BR, breadth of rostrum
BZP, breadth of the zygomatic plate
DAB, depth of the auditory bulla
IOB, interorbital breadth
LBP, length of bony palate
LD, length of diastema
LIF, length of the incisive foramina
LM1-3, coronal length of maxillary toothrow
LR, length of rostrum
ONL, occiptonasal length
PPB, posterior breadth of the bony palate
PPL, postpalatal length
WM1, width of the first upper molar
ZB, zygomatic breadth

Standard descriptive statistics (mean, range,

and standard deviation) were derived for adult
specimens in each species sample. We define
"adult" as the age cohort consisting of animals
that lack the finer juvenile pelage and that possess
fully erupted, though sometimes little worn, third
molars. The mammae formula is presented as the
number of paired postaxial, abdominal, or ingui-
nal teats (sensu Voss & Carleton, 1993).

Where sample sizes permitted, two-sample t-
tests and one-way analyses of variance were ap-
plied to the mensural variables with sex as the
categorical variable. Principal components (PCs)
and canonical variates (CVs) were extracted from
the variance-covariance matrix and computed us-
ing natural logarithmic transformations of the 18
craniodental variables. Loadings are expressed as
Pearson product-moment correlation coefficients
between the PCs or CVs and the original skeletal
and dental variables. Mahalanobis distances be-
tween group centroids were clustered based on the
unweighted pair-group method using arithmetic
averages (UPGMA). All univariate and multivar-
iate computations were generated using Systat
(version 8.0, 1998), a series of statistical routines
programmed for microcomputers.

Taxonomic Comments

Variation Within Eliurus majori and the
Status of E. penicillatus

Within the Marojejy series that we report here
as Eliurus majori are several specimens that pos-

CARLETON & GOODMAN: RODENTS

235

co
o

>
O

-4

-2

-10 12 3
CV 1 (60.7 %)

80

60

40
Mahalanobis D'

20

Montagne d'Ambre (5)
Marojejy (6)
Anjanaharibe-Sud (2)
Andringitra (3)
Ivohibe (4)
Andohahela (1)
Ampitambe (7)

236

FIELDIANA: ZOOLOGY

sess a white-tipped tail. Where specifically noted
in Goodman's field catalogue or verifiable in pre-
served specimens, the terminal section in almost
half of the series (9 of 20) is invested with white
hairs, the white portion ranging from 9 to 54 mm
long and averaging 29 mm. The remainder of the
Marojejy series, and all E. majori from nearby RS
d'Anjanaharibe-Sud, have a tail that is uniformly
dark to the tip, conforming to the diagnosis of E.
majori (Thomas, 1895). In other aspects of their
cranium, dentition, pelage, and size, these nine in-
dividuals resemble typical E. majori as now
known by populations reported elsewhere in the
eastern forest (Goodman & Carleton, 1996, 1998;
Goodman et al., 1997; Goodman, Carleton, &
Pidgeon, 1999).

Possession of a white caudal tip is central to
the diagnosis of the species E. penicillatus, a form
that otherwise recalls a somewhat smallish ver-
sion of E. majori (Thomas, 1908; Carleton, 1994).
Of the eight species of Eliurus he recognized,
Carleton (1994) regarded the specific separation
of E. penicillatus Thomas (1908) from E. majori
Thomas (1895) as the most weakly defended, but
retained them as distinct based on the constancy
of the contrast in tail color and the inadequacy of
sample sizes to convincingly address the problem
(5 specimens of E. majori and 17 of E. penicil-
latus). The variable presence of white tail tips in
the Marojejy series of E. majori invites reevalu-
ation of the status of E. penicillatus, a form so far
recorded only from its type locality, Ampitambe,
in east-central Madagascar.

The vastly improved population samples of E.
majori, now well represented from Montagne
d'Ambre in the north (Goodman et al., 1997) to
Andohahela in the south (Goodman, Carleton, &
Pidgeon, 1999), enhance the geographic scope of
our reappraisal. For multivariate comparisons,
we measured 95 specimens with intact skulls
that represent six general population samples
(operational taxonomic units, OTUs) of E. ma-
jori and one of E. penicillatus, as follows (spe-
cific locality data and museum numbers listed in
Appendix 12-1): (1) RNI d' Andohahela (N = 4);
(2) RS d'Anjanaharibe-Sud (N = 10); (3) RNI

d'Andringitra (N = 17); (4) RS dTvohibe (N =
4); (5) PN de la Montange d'Ambre (N = 25);
(6) PN de Marojejy (N = 17); (7) Ampitambe (N
= 16; type locality of E. penicillatus). The type
specimens of E. majori (BMNH 97.9.1.147, Am-
bohimitombo) and E. penicillatus (BMNH
97.9.1.149, Ampitambe) were entered as un-
knowns for a posteriori classification by the mul-
tipliers generated from seven-group discriminant
function analysis.

Three nonoverlapping clusters are evident in
projections of specimen scores onto the first two
CVs derived (cumulative variance explained =
81.5%). The largest includes those samples of E.
majori from throughout the northern, central, and
southern highlands (OTUs 1-4, 6; Fig. 12-1).
Marginal to these is the series from the isolated
peak in the north, Montagne d'Ambre (OTU 5).
The series from Ampitambe (OTU 7), the type
locality of E. penicillatus, exhibits the strongest
differention among the geographic samples,
which on average are separated from the other
centroids by a generalized distance of 73.5 (Fig.
12-1). Except for the extreme separation of the
Ampitambe series, Mahalanobis distances be-
tween centroids loosely approximate geographic
distances between localities namely, the pair-
group association between Andringitra and Ivo-
hibe on the Central High Plateau, and the linkage
of samples from the northern highlands (Anjana-
haribe-Sud and Marojejy), southern highlands
(Andohahela), and Montagne d'Ambre at succes-
sively greater values. The central geographic lo-
cation of Ampitambe, if correctly interpreted by
Carleton and Schmidt (1990), and the peripheral
phenetic divergence of E. penicillatus depart from
this pattern. Ampitambe, an important Forsyth
Major collecting site and the source of numerous
holotypes of Malagasy mammals, is believed to
be situated along the east-central flank (ca. 900
m) of the Central High Plateau, nearest Andrin-
gitra and Ivohibe in the southern Central High
Plateau among the samples considered here.

Size, as a general factor, is negatively associ-
ated with the first CV, most of the variable cor-
relations being moderately large (about -.60 to

Fig. 12-1. Results of discriminant function analysis performed on 18 log-transformed craniodental variables, as
measured on 95 specimens representing seven OTUs of the Eliurus majori complex. Top, Projection of individual
scores onto the first two canonical variates extracted; polygons enclose the maximal dispersion of specimen scores
around a group centroid, and crosses signify the type specimens of E. majori and E. penicillatus. Bottom, Phenogram
produced from clustering (UPGMA) of Mahalanobis' distances among centroids of the seven OTUs. See Table
12-2.

CARLETON & GOODMAN: RODENTS

237

Table 12-2. Discriminant function results and one-
way ANOVAs derived from analysis of 18 log-trans-
formed craniodental dimensions as measured on 95 adult
specimens representing seven samples of the Eliurus
majori complex.

Correlations

Variable

CV 1

CV2

F (OTU)

ONL

-0.67

0.12

10.8**

ZB

-0.75

0.01

12.6**

BBC

-0.70

-0.13

18.3**

IOB

-0.32

0.39

4.2**

LR

-0.62

0.09

g 7**

BR

-0.53

0.09

7.5**

PPL

-0.59

0.16

7.5**

LBP

-0.64

-0.18

10.3**

LIF

-0.70

0.11

14.0**

BIF

-0.24

0.61

11.5**

LD

-0.61

-0.17

8.5**

BMls

-0.77

0.06

15.3**

PPB

-0.82

0.04

18.7**

DAB

-0.35

0.62

10.6**

BZP

-0.75

-0.23

17.2**

BOC

-0.41

-0.24

4.0*

LM1-3

-0.86

0.21

35.6**

WM1

-0.80

0.15

16.6**

Canonical

correlation

0.95

0.87

Eigenvalue

9.34

3.21

% Variance

60.7

20.8

* P < 0.01; ** P < 0.001.

Abbreviations are explained in the Materials and
Methods section. See also Figure 12-1.

-.80) and nearly all loading highly significantly
on this factor (Table 12-2). Dispersion of OTUs
along CV1 sensibly mirrors one's visual impres-
sion of size differences when reviewing samples,
from the smaller animals of Ampitambe (OTU 7)
and Montagne d'Ambre (OTU 5) to the larger
specimens representing E. majori sensu stricto
(OTUs 1-4, 6). Although nearly as diminutive as
the Ampitambe series in many craniodental mea-
surements, notably size of molars and occipiton-
asal length (Table 12-3), the sample from Mon-
tagne d'Ambre conforms more closely to the E.
majori cluster in certain features of shape. Size of
bullae and breadth of the incisive foramina, which
correlate moderately strongly and positively with
CV2, largely account for this association and the
greater isolation of the topotypic sample of E.
penicillatus (Table 12-2, Fig. 12-1). The first two
PCs extracted from ordination of only the 41
specimens representing the Montagne d'Ambre
and Ampitambe OTUs (not illustrated) disclose
no overlap of the two series and emphasize the
same shape contrasts on PCII (% variance ex-

plained = 43.5% and 23.7% on PCI and PCII,
respectively), in addition to others whose signifi-
cance is masked in the full data set (the shorter
bony palate and narrower zygomatic plate of E.
penicillatus see Table 12-3).

The pattern of divergence in canonical space
logically corresponds to jackknifed classification
iterations, which commonly produced specimen
misassociations among the large cluster of five
OTUs (% correct = 25-76), uncommonly for that
from Montagne d'Ambre (% correct = 92), and
never for that from Ampitambe (% correct =
100). The holotype of E. penicillatus was as-
signed (P = 1 .00) to OTU 7 from Ampitambe, its
topotypic complement; that of E. majori was clas-
sified (P = 0.82) with OTU 3 from Andringitra,
the locality geographically nearest to Ambohimi-
tombo, its type locality.

In summary, we regard these multivariate pat-
terns of craniodental differentiation as insufficient
to justify the synonymy of E. penicillatus Thomas
(1908) under E. majori Thomas (1895). Other in-
formation sources must be brought to bear on the
issue, as well as the collection of new material
from or near Ampitambe, the type locality of Ma-
jor's penicillatus. The substantial divergence re-
vealed for the sample from Montagne d'Ambre
also merits further evaluation, but in craniodental
proportions, the affinity of this isolated population
lies with E. majori proper. Relationships inferred
from cytochrome b data actually affiliate the
Montagne d'Ambre sample more closely with
other E. majori from the northern highlands (An-
janaharibe-Sud) than with those from central and
southern Madagascar (Jansa, 1998).

Variation Within Eliurus webbi Apropos the
E. majori Complex

Evaluation of craniodental variation among
populations of Eliurus webbi is instructive in light
of the patterns revealed for the E. majori complex
(that is, including E. penicillatus). The distribu-
tions of both forms span the full extent of eastern
humid forest, as reported from the PN de la Mon-
tagne d'Ambre in the far north to parcel 1 of the
RNI d'Andohahela in the far south (Goodman &
Carleton, 1996, 1998; Goodman et al., 1996;
Goodman, Carleton, & Pidgeon, 1999). Although
geographic boundaries are broadly congruent,
their elevational ranges are superpositioned, sug-
gesting contiguous allopatry within the eastern
humid forest biome. Eliurus webbi occurs in low-

238

FIELDIANA: ZOOLOGY

Table 12-3. Comparison of craniodental measurements of adult Eliurus majori and E. penicillatus from localities
in eastern Madagascar.

E. majori

E. penicillatus

Variable

Mt. d'Ambre

(n = 27)

Marojejy

(n = 17)

Andringitra

(n = 17)

Ampitambe
(n = 16)

ONL

36.5 1.2
33.5-38.1

39.1 1.6
35.9-41.3

37.6 1.0
35.5-39.4

36.1 1.3
32.9-37.4

ZB

18.5 0.7
16.9-19.5

19.9 0.7
18.9-20.8

19.1 0.5
18.1-19.8

17.9 0.9
15.9-19.2

BBC

14.1 0.4
13.2-14.8

15.1 0.5
14.4-15.9

14.3 0.4
13.4-14.8

13.6 0.4
12.7-14.1

IOB

5.2 0.2
4.8-5.6

5.5 0.3
5.1-6.1

5.3 0.2
4.9-5.7

5.3 0.1
5.1-5.6

LR

12.6 0.6
11.3-13.5

13.5 0.6
12.2-14.3

13.1 0.5
11.6-14.0

12.4 0.6
10.8-13.2

BR

7.0 0.3
6.3-7.6

7.5 0.4
6.6-8.1

7.1 0.2
6.8-7.6

6.9 0.4
6.1-7.4

PPL

12.9 0.6
11.4-13.8

13.8 0.5
12.8-14.3

13.4 0.5
12.4-14.6

12.9 0.6
11.5-13.8

LBP

7.4 0.4
6.6-8.5

7.9 0.6
6.7-8.7

7.5 0.4
6.9-8.6

6.8 0.3
6.3-7.4

LIF

5.5 0.4
4.7-6.3

6.1 0.4
5.5-7.0

6.3 0.3
5.6-6.7

5.5 0.3
5.0-5.9

BIF

2.3 0.2
2.0-2.6

2.6 0.1
2.3-2.8

2.7 0.2
2.4-3.1

2.7 0.2
2.4-2.9

LD

10.5 0.5
9.2-11.3

11.0 0.6
9.9-12.2

10.8 0.4
10.0-11.8

9.8 0.6
8.5-10.7

BM1S

7.5 0.3
7.1-8.3

8.2 0.3
7.6-8.5

8.1 0.3
7.1-8.6

7.3 0.3
6.8-7.9

DAB

5.0 0.2

4.7-5.4

5.5 0.2
5.3-5.9

5.2 0.1
5.1-5.5

5.4 0.2
5.0-5.6

BZP

3.3 0.2
2.9-3.6

3.8 0.3
3.3-4.2

3.5 0.2
2.9-3.9

2.9 0.3
2.5-3.4

BOC

8.3 0.3
7.7-8.7

8.4 0.3
8.0-8.9

8.5 0.3
8.0-9.0

8.0 0.3
7.4-8.4

LM1-3

6.07 0.21
5.69-6.43

6.69 0.23
6.21-7.10

6.65 0.24
6.13-7.15

6.03 0.15
5.74-6.27

WM1

1.61 0.09
1.43-1.79

1.78 0.07
1.65-1.91

1.76 0.07
1.66-1.88

1.58 0.05
1.49-1.68

Abbreviations are explained in the Materials and Methods section. Sample parameters are
deviation, and range.

given as mean standard

land rain forest, usually below 875 m, whereas E.
majori inhabits montane and sclerophyllous mon-
tane forest, usually above 1200 m (see Discus-
sion, below). To date, the two species have been
documented syntopically only on Montagne
d'Ambre at 1000 m (Goodman et al., 1997).

As in the case of E. majori, availability of ma-
terial has much improved the study of E. webbi.
Ninety specimens representing nine general local-
ities were measured for morphometric compari-

sons (see Appendix 12-1): (1) RNI d'Andohahela
(N = 7); (2) RS d'Anjanaharibe-Sud (N = 6); (3)
RNI d' Andringitra (N = 19); (4) 20 mi S Fara-
fangana (N = 12; type locality of E. webbi); (5)
RS d'lvohibe (N = 7); (6) PN de la Montagne
d'Ambre (N - 6); (7) PN de Marojejy (N = 5);
(8) NE lowlands in the vicinity of Antongil Bay
(N = 8); and (9) PN de Ranomafana (N = 20).
As before, the type specimen of E. webbi (BMNH
47.1576) was entered as an unknown for a pos-

CARLETON & GOODMAN: RODENTS

239

CD

i
C\J
>

2 -

1 -

-

-2 -

-3 -

-4

40

-1 1

CV 1 (39.6 %)

Montagne d'Ambre (6)
Anjanaharibe (2)
NE Lowlands (8)
Ranomafana (9)
Ivohibe (5)
Andringitra (3)
Marojejy (7)
Farafangana (4)
Andohahela (1)

30

20

10

Mahalanobis D

240

FIELDIANA: ZOOLOGY

teriori classification by the multipliers generated
from nine-group discriminant function analysis.

No clearly discrete, nonoverlapping clusters of
OTUs were evident in projections of specimen
scores onto the first two CVs derived (compare
Fig. 12-2 with 12-1, and Table 12-4 with 12-2).
The cumulative variance explained is smaller than
in the example of E. majori (59.2% vs 81.5%), as
are other multivariate statistics (canonical corre-
lations, eigenvalues) that convey the success of
distilling covariation among the original variables
into fewer dimensions. Except for the unexpected
pair-group association of Farafangana (southeast-
ern) and Marojejy (northeastern), UPGMA clus-
ters coarsely reflect geographic regions and rela-
tive isolation namely, the union of Ranomafana,
Ivohibe, and Andringitra along the Central High
Plateau, the junction of Anjanaharibe-Sud and the
lowlands OTU in the northeast, and the incremen-
tally greater distances linking Andohahela from
the southern Anosyenne Mountains and the sam-
ple on Montagne d'Ambre. On average, however,
an OTU recovered only one-third of its preas-
signed specimens in post hoc jackknifed classifi-
cations by the discriminant coefficients generated
(% correct = 0-57). Although the holotype of E.
webbi falls within the maximal spread of Farafan-
gana scores (OTU 4), the shorter distance between
it and the centroid of OTU 7 (Marojejy) instead
resulted in its classification with that sample, at a
low probability level (P = 0.45).

We view these multivariate patterns of cranio-
dental variation as indicative of a single, widely
distributed species, E. webbi Ellerman (1949).
These numerical results recall the amount of dif-
ferentiation that Carleton and Goodman (1998)
encountered among geographic samples of E. tan-
ala, a form that occurs over nearly the same geo-
graphic extent as E. webbi but more broadly over-
laps the latter's altitudinal range (sympatric from
450 to 875 m, as so far known). Although Carle-
ton and Goodman construed the intraspecific var-
iation of E. tanala as a weak north-south size
cline, the variable correlation coefficients and
sample statistics derived for E. webbi obscure any
simple clinal explanation. As for E. majori proper,

Table 12-4. Discriminant function results and one-
way ANOVAs derived from analysis of 18 log-trans-
formed craniodental dimensions as measured in 90 adult
specimens representing nine samples of Eliurus webbi.

Correlations

Variable

CV 1

CV2

F (OTU)

ONL

0.25

0.31

3.3**

ZB

0.23

0.17

2 8**

BBC

0.43

0.35

4.8***

IOB

0.33

0.04

2.3*

LR

0.16

0.10

1.7

BR

0.16

0.22

2.1*

PPL

0.14

0.45

3.5**

LBP

0.29

0.07

3.7**

LIF

-0.18

0.47

2.6*

BIF

0.61

-0.09

1 g***

LD

0.18

0.21

1.8

BMls

0.18

0.14

1.5

PPB

0.14

0.24

2.4*

DAB

0.47

0.41

1 4***

BZP

0.55

0.37

7.8***

BOC

0.14

0.08

1.9

LM1-3

0.21

-0.03

2.6*

WM1

-0.20

0.30

2.8**

Canonical

correlation

0.87

0.78

Eigenvalue

3.07

1.52

% Variance

39.6

19.6

* P < 0.05; ** P < 0.01; *** P < 0.001.
Abbreviations are explained in the Materials and
Methods section. See also Figure 12-2.

the sample of E. webbi from Montagne d'Ambre
exhibits the strongest craniodental differentiation,
although of lesser magnitude (Fig. 12-2). In ad-
dition to smaller size, ventral pelage color of the
Montagne d'Ambre series is exceptional for the
expansive patches of creamy-white hairs, in con-
trast to the uniformly dingy gray underparts typ-
ical of E. webbi (Carleton, 1994).

Compared to the results obtained for the low-
land species E. webbi, the greater structure ap-
parent in the data matrix of the E. majori complex
plausibly follows from the increased likelihood of
populational fragmentation of a montane-dwelling
organism and subsequent differention among the
geographic isolates. Palynological data and distri-

Fig. 12-2. Results of discriminant function analysis performed on 18 log-transformed craniodental variables, as
measured on 90 specimens representing nine OTUs of Eliurus webbi. Top, Projection of individual scores onto the
first two canonical variates extracted; polygons enclose the maximal dispersion of specimen scores around a group
centroid, and the cross signifies the type specimen of E. webbi. Bottom, Phenogram produced from clustering
(UPGMA) of Mahalanobis' distances among centroids of the nine OTUs. See Table 12-4 and compare with Figure
12-1.

CARLETON & GOODMAN: RODENTS

241

Table 12-5. External measurements and sample statistics for adult rodents collected in the PN de Marojejy.

Species

TOTL

HBL

TL

HFL

EL

WT

Rattus rattus
Eliurus grandidieri

Eliurus majori

Eliurus minor
Eliurus tanala
Eliurus webbi

Gymnuromys roberti
Voalavo gymnocaudus

392, 430

160. 180

224, 224

33.. 33

25. 27

83.5, 145

291.3

124.8

161.5

28.3

20.1

50.6

10.0

6.6

8.7

1.0

0.9

4.0

275-309

111-164

144-176

26-31

19-23

42.0-62.0

(n = 42)

(n = 59)

(n = 42)

(n = 59)

(n = 60)

(n = 59)

363.9

161.9

189.5

29.7

18.6

111.8

21.7

8.2

15.1

0.9

1.0

16.1

321-394

146-175

164-213

28-31

17-20

89.0-151

(n = 20)

(n = 23)

(n = 20)

(n = 23)

(n = 23)

(n = 22)

239, 245

107, 113

126, 129

21. 23

16. 18

37.5. 50.5

359

158, 162

140

30, 34

22, 24

102. 105

324.8

142.5

176.5

29.5

22.4

74.5

14.3

5.5

8.8

1.5

1.1

9.7

303-344

135-152

161-186

27-31

21-24

60.5-90.5

(n = 6)

(n = 11)

(n = 6)

(n = 10)

(n = 10)

(n = 11)

395

175

206

38

23

162

211.2

84.6

120.6

18.8

15.0

22.2

8.3

4.2

5.3

1.1

0.0

3.3

200-219

80-90

113-126

17-20

15-15

17.0-25.5

(n = 5)

(n = 5)

(n = 5)

(n = 5)

(n = 5)

(n = 5)

Abbreviations are explained in the Materials and Methods section. Sample parameters are given as mean
deviation, range, and number.

standard

butional information gathered on other Malagasy
vertebrates certainly establish the profound verti-
cal changes experienced by montane communities
throughout the Late Pleistocene and into the Ho-
locene (Burney, 1987, 1997; Gasse et al., 1994;
Raxworthy & Nussbaum, 1996). The challenge is
to interpret this data structure in light of current
taxonomy and defensible criteria for species rec-
ognition, which in the case at hand persuades us
to continue to regard E. penicillatus as distinct
from E. majori.

Accounts of Species

General observations, information on the nat-
ural history, and the elevational range of each ro-
dent species captured during the inventory of the
PN de Marojejy are presented under the subhead-
ings Distribution, Ecology and Reproduction,
Comments (if appropriate), and Specimens Ex-
amined. The exact position and coordinates of the
sites presented under this last subheading can be
found in Chapter 1 . More detailed criteria for spe-
cies identification and discussion of alpha-level
taxonomic problems are presented in Carleton

(1994), Carleton and Goodman (1996, 1998),
Goodman and Carleton (1996, 1998), and Good-
man, Carleton, & Pidgeon (1999). External mea-
surements and masses are given herein for the ro-
dent species captured during the 1996 survey to
aid researchers in identifying the rodents of north-
ern Madagascar and to provide baseline data used
in some analyses presented in the Discussion (Ta-
ble 12-5).

Family Muridae: Subfamily Murinae

Rattus rattus (Linneaus, 1758)

Distribution This introduced rodent has a
broad distribution on the island. It is found as a
commensal near human habitations and agricul-
tural areas, in disturbed forest habitats, and in pri-
mary forest. A few Rattus rattus were trapped at
1250 and 1625 m. totaling only 4 (4.8%) of the
83 rodents captured at those middle elevations
(Table 12-6) and suggesting that the species is un-
common within the PN de Marojejy.

Ecology and Reproduction On the basis of
a limited number of captures of this species in the

242

FIELDLANA: ZOOLOGY

Table 12-6. Number of individuals captured in live traps for each species of small mammal at all five elevations
surveyed in the PN de Marojejy.

Species or
parameter
measured

450 m
(690)

775 m
(800)

1250 m

(700)

1625 m
(600)

1875 m

(600)

Tenrecinae

Microgale cowani
Microgale gracilis
Microgale talazaci
Oryzorictes hova*

1

4

2

1
1

1

Murinae

Rattus rattus

3

1

Nesomyinae
Eliurus grandidieri
Eliurus majori
Eliurus minor
Eliurus tanala
Eliurus webbi
Gymnuromys roberti
Voalavo gymnocaudus

9

2
2
4

44
2

1

1

11
19

1

8

1

Total no. of individuals
Trap success, %
Total no. of rodents trapped
Total no. of rodent species
Rodent trap success, %

10
1.4
9
1
1.3

12
1.5
8
3
1.0

53

7.6
51

5

7.3

34

5.7
32

4

5.3

10
1.6
9
2
1.5

Total no. of nesomyines trapped
Total no. of nesomyine species
Nesomyine trap success, %

9

1
1.3

8
3
1.0

48
3
6.9

31

2
5.2

9
2
1.5

* We follow Goodman, Jenkins, and Pidgeon (1999) in considering O. talpoides a synonym of O. hova.
Total number of trap-days accrued is given in parentheses. Pitfall results are excluded.

1250 to 1625 m zones of the PN de Marojejy, trap
sets that yielded this animal were equally divided
between ground and arboreal trap placements (Ta-
ble 12-7). One of the successful sets off the
ground was on a large, horizontal, moist, and pre-
sumably slippery branch of a tree that was cov-
ered with a dense epiphytic growth.

Both male Rattus rattus obtained in the reserve
had scrotal testes. Of the females obtained, one
had large mammae and a perforated vagina and
the second was a subadult with small mammae
and an imperforate vagina. The mammae formula
of the adult female was 1-1-2.

Comments Remarkably few Rattus rattus
were captured on the Marojejy Massif as com-
pared to the Anjanaharibe-Sud Massif. In the
1 250 m zone of the former locality, three individ-
uals were trapped in 700 trap-nights, and in the
1625 m zone one individual was trapped in 600
trap-nights. In the RS d' Anjanaharibe-Sud, this
species was found across the complete transect,
from 875 to 1950 m, and was most common in
the 1550 m zone, where 13 individuals were cap-

tured in 500 trap-nights (Goodman & Carleton,
1998). On the basis of short-term field studies at
these two sites (which are only a few tens of ki-
lometers distant) conducted two years apart dur-
ing the same season, no clear explanation can be
offered for the differences in elevational distri-
bution and relative densities of this species.

In spite of the now substantial elevational data
for Rattus rattus on various forested mountains in
eastern Madagascar, a general pattern that ex-
plains the species' distribution cannot be gleaned.
The highest density recorded to date is in the for-
est of the PN de la Montagne d'Ambre, in the
extreme north, where 103 R. rattus were captured
at 1350 m in 750 trap-nights (Goodman et al.,
1996). In a broad swath of other elevational
zones, such as Marojejy and Andohahela (Good-
man, Carleton, & Pidgeon, 1999), few individuals
of this species were captured in a nearly compa-
rable number of trap-nights. Regression analyses
were conducted to assess geographic location and
variation in Rattus population density found at PN
de la Montagne d'Ambre (12.5S, Goodman et al.,

CARLETON & GOODMAN: RODENTS

243

-

51

z

a. ?

5? >-

3 **

=

X

=

5 S

u

A

U

G

>

E

s _
3 2

. .-SB

** a *>

5 =

fig g

**

$1

< M

3
Si

f-

3 S

-

ir, rN <N

O r*-, <N \C

C* c*".

3 CI </-.

O 3C

(N

O CN (N fN

C. (N C". C ~

X C ^ N C it. (S (N vC

<n r- r ~ r-

see E

? s

ill! s

1 1-| 7 1| ill i?

"5-3 S" 55 -2 -3 -3 -2 -2

244

FIELDIANA: ZOOLOGY

s

fj

Z E

is
3 **
09

S o

3 Ja 7 1

-J *- a

- s
. - -

J Sio

CO |

111

111

;-

4*2

.2 *2
33 < oc
"5
5
a

a

o I

i m

9 .2*

- a
>

es

"Si

1996), RS d'Anjanaharibe-Sud (14S, Goodman
& Carleton, 1998), Marojejy (14.5S, this chap-
ter), RNI d'Andringitra (22S, Goodman & Carle-
ton, 1996), and RNI d'Andohahela (24.5S, Good-
man, Carleton, & Pidgeon, 1999). No significant
relationship was demonstrable between the lati-
tude of these mountains and that elevational zone
with the highest density of Rattus (R 2 = 0.32, df
= 4, F = 1.42, P = 0.32), or between latitude
and number of captures in the zone of highest
density as measured by trap success (R 2 = 0.12,
df= 4, F = 0.41, P = 0.57).

Specimens Examined 1 1 km NW Mananteni-
na, Antranohofa, 1250 m (FMNH 159649; UA-
SMG 8362, 8399); 10.5 km NW Manantenina,
1625 m (FMNH 159650).

Family Muridae: Subfamily
Nesomyinae

Eliurus grandidieri Carleton and Goodman,
1998

Distribution The geographic range of E.
grandidieri, as so far understood, includes the
mountains surrounding the Andapa Basin in the
north, specifically the Anjanaharibe-Sud and Ma-
rojejy massifs, and the highlands at least as far
south as the Andranomay Forest of the Anjozo-
robe district (Carleton & Goodman, 1998; Good-
man et al., 1998). A straight-line distance of about
340 km separates Andranomay and the mountains
surrounding the Andapa Basin. The species is
known over an elevational range of 1250-1875 m
in the mountains surrounding Andapa and from
about 1300 m in the Andranomay Forest. Little
field survey of small mammals has been con-
ducted in the area between these two locales, par-
ticularly on rodents of upland forests, but E. gran-
didieri may be anticipated in appropriate habitat
across the region.

Ecology and Reproduction In general, this
species is terrestrial, although a small percentage
of individuals were trapped in sets placed off the
ground (Table 12-7). Documented at 1250, 1625,
and 1875 m, at least half of the E. grandidieri
obtained on the ground were captured among
roots of standing trees, by fallen tree trunks, or
under rocks. Such sets were frequently placed in
front of hollows and tunnels, suggesting that the
species occupies subterranean burrows. It was

CARLETON & GOODMAN: RODENTS

245

fc- . a. Z

C3 u "O

<N(N0C(N<N0O <N ir-j

(N C"i O ON

5; O
^r

m o

-"JMO

r<l c~, o

IT!

(N

<N <N C-,
<N

(N (N (N

~5j ^ ^

5 S 3* -5 2

>- *- - n -*

3:s

^

rr

3

E c

cs

CS <U

#

60

ON

<D O

soc.

r~

-s =

r~

._ <D

(N

o

^ s

^ *>g S 5 * ^

a 2 ? ? ? ? II

s .a .3 .a .s .a 5 3

c S3 S3 S3 S3 S3 o'^

^

en

(N

>,

JO

E

0)

c

00

=

M

'>,

c

E

'c_

9

Q.

tu

-->

u

e/5

o

3

>,

o

-

E

Cfl

85

C

3

.t^

3

=E

-o

as

u

3

=

E

***

ii

2^

v:

._

u

t/5

S3

3

71

u

II

3

<

u

a

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u

c:

r.

X)

h

<

cd

246

FIELDIANA: ZOOLOGY

rarely captured in areas of dense leaf litter or open
understory within the forest.

Arboreal sets that produced samples of E. gran-
didieri were exclusively positioned on limbs and
trunks >10 cm in diameter (Table 12-7). The spe-
cies apparently prefers larger limb and trunk sub-
strates when climbing since, within each eleva-
tional zone, at least as many traps were placed on
smaller vines, limbs, and trunks (i.e., <10 cm in
diameter).

Some differences in reproductive activity were
found between the age and sex classes of E. gran-
didieri in the three elevational zones where it was
captured (Table 12-8). In the 1250 m zone, about
two-thirds of the individuals captured were adults.
Of 21 males, 14 had scrotal or nearly scrotal testes
and the balance had abdominal testes; of 20 fe-
males, 12 had large mammae and perforated va-
ginas and eight possessed small mammae and im-
perforate vaginas. At 1625 m, all seven males ob-
tained had scrotal testes; one of four females had
enlarged mammae and a perforated vagina; the
other three females were subadults with imper-
forate vaginas. Finally, in the 1875 m zone, one
of four males trapped had scrotal testes and the
other three possessed abdominal testes; the four
females were evenly split between individuals
with large mammae and perforated vaginas and
those with small mammae and imperforate vagi-
nas.

All female E. grandidieri captured possessed
three pairs of mammae (1-1-1, n = 26), a number
and pattern identical to that found for this species
in the RS d'Anjanaharibe-Sud (Goodman &
Carleton, 1998). Although many females pos-
sessed enlarged mammae, only one of 28 ap-
peared to be actively lactating. Furthermore, of
seven females examined, not one contained em-
bryos, and only one had placental scars, in this
instance three. Reproduction in this species may
be seasonal; the October-November 1996 survey
of the PN de Marojejy was conducted after the
most recent breeding period. A pattern of seasonal
reproduction appears to hold for this species in
the RS d'Anjanaharibe-Sud, where the survey was
conducted between late October and November
1994 (Goodman & Carleton, 1998).

Comments Eliurus grandidieri was abundant
in the 1250 m zone, where 44 individuals were
captured in a trap line with 100 different trap sta-
tions (Table 12-6). Along a 120 m stretch of linear
trap line, 18 E. grandidieri were captured, yield-
ing a density of one individual per 6.6 m. In spite
of this spatial clumping, the occurrence of multi-

ple captures in a single trap does not significantly
deviate from a Poisson distribution. Of the traps
that yielded this species, 21 captured single indi-
viduals, seven captured two individuals, and three
trapped three individuals.

The fine sample of E. grandidieri from the PN
de Marojejy conforms closely to the morpholog-
ical description of the type series from the RS
d'Anjanaharibe-Sud (Carleton & Goodman,
1998), as might be expected from their geographic
proximity. Tail length, as noted for the species, is
relatively long on the Marojejy specimens (TL
129% of HB, Table 12-6), and where recorded or
observable on preserved material, the caudal tip
consistently possesses white hairs, the white ter-
minal section 3-33 mm in length and averaging
21.3 mm (N = 9). An alisphenoid strut is char-
acteristically present and well developed on Ma-
rojejy crania (present on both sides in 22 skulls,
present on one side in three, and absent in none),
whereas the strut's occurrence is more variable in
the type series (absent on both sides in four of 14
skulls).

Specimens Examined 1 1 km NW Mananteni-
na, Antranohofa, 1250 m (FMNH 159579-
159609, 159698-159703; UA-SMG 8342, 8347,
8361, 8386); 10.5 km NW Manantenina, 1625 m
(FMNH 159610-159618, 159704, 159705); 11
km NW Manantenina, at source of Andranomi-
fototra River, 1875 m (FMNH 159619-159623,
159706-159708).

Eliurus majori Thomas, 1895

Distribution The anticipated presence of E.
majori in montane and sclerophyllous montane
vegetation (1250-1875 m) of the PN de Marojejy
conforms to our broad understanding of its distri-
bution, documented in similar settings from Mon-
tange d'Ambre in the north to the Anosyenne
Mountains in the south.

Ecology and Reproduction Approximately
two-thirds of all E. majori captured had entered
live traps placed aboveground (Table 12-7). Of
the 15 individuals trapped in arboreal sets, 11
were obtained on limbs and trees >10 cm in di-
ameter. These trapping results suggest that the
species in the PN de Marojejy is largely scanso-
rial, preferring thicker arboreal substrates, a gen-
eralization that seems to hold across its range
(e.g., Goodman & Carleton, 1996, 1998).

Most examples of E. majori (20) were recorded
in the 1625 m zone, 19 of these obtained in a

CARLETON & GOODMAN: RODENTS

247

linear trap line extending 385 m. Eight (42%) of
19 were captured along a 23 m section of the line,
consisting of six consecutive traps, two of which
yielded two individuals each and one three indi-
viduals. Other E. majori trapped within this ele-
vational zone did not show such a clumped dis-
tribution.

Signs of breeding are evident within each ele-
vational zone. At 1625 m, 11 of 12 males ob-
tained were either fully or partially scrotal. Of
seven females obtained with data on reproductive
condition, five had enlarged mammae but none
showed signs of active lactation, and only one
showed any sign of estrus her uterine walls were
highly vascularized. The mammae formula is con-
sistently 1-1-1 (n = 7). Only two individuals were
trapped at 1250 m and a single individual at 1875
m (Table 12-6), all of them adult males with scro-
tal testes.

Specimens Examined 1 1 km NW Mananteni-
na, Antranohofa, 1250 m (FMNH 159624,
159709); 10.5 km NW Manantenina, 1625 m
(FMNH 159625-159637, 159710-159715; UA-
SMG 8463); 1 1 km NW Manantenina, at source
of Andranomifototra River, 1875 m (FMNH
159638).

Eliurus minor Major, 1896

Distribution Eliurus minor, one of the most
widely distributed nesomyine rodents, occurs
throughout the eastern humid forest, from Mon-
tagne d'Ambre in the north to the Tolagnaro re-
gion in the south (Goodman et al., 1996; Good-
man, Carleton, & Pidgeon, 1999). In the PN de
Marojejy, the species was documented over mid-
dle elevations, 775-1625 m; elsewhere the species
has been recorded from near sea level (vicinity of
Antongil Bay) to about 1 800 m (west of Andapa)
(Carleton, 1994).

Ecology and Reproduction Using live traps,
two E. minor were captured at the 775 m site and
a single animal at 1250 m. Single individuals were
taken in pitfall buckets in the 775 m and 1625 m
zones (see Chapter 11); both were found during
dawn checks after heavy rain the previous night.

Of the three specimens obtained in live traps,
two were taken off the ground and one on the
ground (Table 12-7). Both arboreal sets were at a
height of 1.5-2 m on horizontal saplings or
branches <4 cm in diameter. At other sites, the
species has been taken on a greater range of sub-
strate diameters, including on branches and trunks

>10 cm in width (Goodman & Carleton, 1998;
Goodman, Carleton, & Pidgeon, 1999).

No external sign of breeding was noted in the
two females captured, although both males col-
lected had scrotal testes (Table 12-8).

Specimens Examined 10 km NW Mananteni-
na, along tributary of Manantenina River, 775 m
(FMNH 159716, 159717); 11 km NW Mananten-
ina, Antranohofa, 1250 m (FMNH 159639; UA-
SMG 8343); 10.5 km NW Manantenina, 1625 m
(FMNH 159640).

Eliurus tanala Major, 1896

Distribution This species also ranges broad-
ly across much of the eastern humid forest, so that
its presence in the PN de Marojejy two individ-
uals obtained in the 775 m zone was anticipated.
On massifs such as Anjanaharibe-Sud, Andringi-
tra, and Andohahela, E. tanala has been trapped
over a broader swath of elevations, typically into
middle montane forest formations as high as 1625
m (Goodman & Carleton, 1996, 1998; Goodman,
Carleton, & Pidgeon, 1999).

Ecology and Reproduction Both individuals
were trapped at arboreal stations, one positioned
at 1 m on a 2-cm-diameter downed tree trunk and
the other at 1.5 m on a 20-cm-diameter fallen tree
trunk that spanned a small stream (Fig. 12-3). In
nearby RS d' Anjanaharibe-Sud, five of the six E.
tanala captured were taken in terrestrial sets
(Goodman & Carleton, 1998). At other sites, cap-
ture of this species seems to be almost evenly di-
vided between ground and arboreal sets (Good-
man & Carleton, 1996; Goodman, Carleton, &
Pidgeon, 1999).

The two E. tanala obtained, both females, were
reproductively active. One had large mammae and
two placental scars (one per each uterine horn), and
the second was lactating and had three placental
scars (two in the right horn, one in the left). The
mammae formula in these individuals is 1-1-1.

Specimens Examined 10 km NW Mananteni-
na, along tributary of Manantenina River, 775 m
(FMNH 159696, 159697).

Eliurus webbi EMerman, 1949

Distribution First described as a localized
subspecies of E. myoxinus (Ellerman, 1949), the
emerging distributional picture of this species re-
veals that it is the most broadly distributed neso-

248

FIELDIANA: ZOOLOGY

Fig. 12-3. View of transitional lowland-montane forest at 775 m. An example of Eliurus tanala was captured
alive in a trap placed on a 20-cm-diameter tree that had fallen across the small stream pictured here (trap placement
category = above ground, on limb or trunk greater than 10 cm in diameter). Other Eliurus found in this kind of
forest include E. minor and E. webbi. Nesomys cf. rufus was observed at this elevation but not captured.

myine of the eastern humid forest. It is now known
to inhabit littoral forests of Tampolo (Rakoton-
dravony et al., 1998) and near Tolagnaro (Good-
man, Carleton, & Pidgeon, 1999), as well as low-

land formations across the full latitudinal expanse
of the humid forest biome (Carleton, 1994; Good-
man & Carleton, 1996, 1998; Goodman et al.,
1998). Consistent with this lower elevational set-

CARLETON & GOODMAN: RODENTS

249

ting. E. webbi was captured only in the 450 and
775 m zones of the PN de Marojejy.

Ecology and Reproduction Of 13 E. webbi
trapped, seven were captured on the ground and
six in arboreal sets (Table 12-7). All individuals
captured on the ground were taken next to rotten
wood, roots, trunks, or rock outcrops. All traps
placed aboveground were on substrates <10 cm
in diameter, as in the RNI d* Andringitra. where
most E. webbi were captured on branches and
vines < 10 cm in diameter (Goodman & Carleton.
1996). In the RS d'Anjanaharibe-Sud and RNI
d'Andohahela. on the other hand, a greater per-
centage of individuals was trapped on thicker sub-
strates (Goodman & Carleton. 1998: Goodman.
Carleton. & Pidgeon. 1999).

The condition of the reproductive organs sug-
gested that few individuals had reached sexual
maturity, the elevational samples composed of
many subadults. In the 450 m zone, one of four
males had scrotal testes and two of five females
had large mammae and perforated vaginas. In the
775 m zone, the testes of one male had slightly
descended and in the other were fully abdominal;
of the three females, one had large mammae, a
perforated vagina, and two placental scars and
two had inconspicuous mammae and imperforate
vaginas. The mammae formula in the three fe-
males examined was 1-1-1.

The timing of reproduction among populations
of E. webbi seems to vary without a discernible
pattern based on the field results so far obtained.
For example, during the survey of the RS
d*Anjanaharibe-Sud. conducted between mid-Oc-
tober and late November 1994. the majority of the
E. webbi captured (875 m) showed signs of on-
going reproductive activity (Goodman & Carle-
ton. 1998). Since the Anjanaharibe-Sud and Ma-
rojejy massifs are at nearly the same latitude, sim-
ple explanations of clinal variation in reproductive
level are plainly insufficient. In like manner, sur-
veys conducted during the same calendar period
in the southern protected areas of Andringitra and
Andohahela (Goodman & Carleton. 1996: Good-
man. Carleton. & Pidgeon. 1999) revealed con-
trasting levels of breeding activity in this species.

Specimens Examined 8 km NW Manantenina.
along tributary of Manantenina River. 450 m
(FMNH 159718-159720. 159641-159645: UA-
SMG 8275): 10 km NW Manantenina. along trib-
utary of Manantenina River. 775 m (FMNH
159646. 159721-159723: UA-SMG 8294).

Gymnuromys roberti Major, 1896

DiSTRiBiTiON-Although infrequently recorded,
this rodent is known to have a broad latitudinal
distribution across the eastern humid forest. With
the exception of Montagne d'Ambre in the ex-
treme north. G. roberti has been captured at ev-
ery eastern humid forest site surveyed by Good-
man, but it appears to be absent at several seem-
ingly appropriate sites on the Central High Pla-
teau, for example in the RS d'Ambohitantely
(Stephenson et al.. 1994; Goodman & Rakoton-
dravony. 2000). Gymnuromys roberti is an elu-
sive terrestrial species of middle elevations, al-
though at some places, such as Andringitra, it
has been trapped at lower elevations (720 m).
Only one individual was captured (1250 m) in
the PN de Marojejy.

Ecology and Reproduction The single spec-
imen, an adult female, was collected in a Sherman
trap placed at a hole opening into a fallen log that
was rotting and covered with epiphytes and moss
(Table 12-7). The individual possessed large
mammae, without embryos but with two placental
scars, one per each horn of the uterus. An Eliurus
grandidieri was obtained in this same trap on the
following night.

Specimens Examined 1 1 km NW Mananteni-
na, Antranohofa. 1250 m (FMNH 159724).

Nesomys cf. rufus Peters, 1870

DiSTRiBUTiON-Although N. rufus is another
ubiquitous inhabitant of the eastern humid forest,
none was captured during the 1996 small mammal
inventory of the PN de Marojejy. This richly col-
ored, large diurnal rodent was one of the most
abundant, commonly observed, and frequently
trapped nesomyines in the nearby RS
d'Anjanaharibe-Sud (Goodman & Carleton.
1998). However, individuals of Nesomys, presum-
ably referable to N rufus. were occasionally ob-
served in the 775 and 1250 m zones of the PN de
Marojejy. Other visual records of the species have
been previously reported from the southwestern
portion of the massif at 800 m and in the north-
west portion between 1000 and 1400 m (Duck-
worth & Rakotondraparany. 1990). The failure to
trap Nesomys during the 1996 survey may relate
to an atypically low population density during the
Marojejy inventory.

Ecology All Nesomys were observed during
the day. mostly in areas of open understory forest

250

FTELDIANA: ZOOLOGY

with relatively dense leaf litter, typical habitat for
this species.

Voalavo gymnocaudus Carleton and Goodman,
1998

DiSTRiBUTiON-This recently described form was
previously known from the RS d'Anjanaharibe-
Sud, at 1300 and 1950 m (Carleton & Goodman,
1998). Accordingly, its presence in the PN de Ma-
rojejy was expected, and individuals of V. gym-
nocaudus were captured between 1260 and 1875
m. These occurrences extend the range of the spe-
cies slightly farther to the east, a distribution so
far localized to mountains ringing the Andapa Ba-
sin.

Ecology and Reproduction Voalavo gym-
nocaudus has been captured on the slopes of the
Marojejy and Anjanaharibe-Sud massifs, between
1260 and 1950 m and in vegetational formations
ranging from montane forest to dwarf upper mon-
tane/sclerophyllous forest. Of the six individuals
obtained in the PN de Marojejy, five were caught
in pitfall buckets (see Chapter 11) and one in a
Sherman trap. The live-trapped individual (1625
m) was collected in a set placed on a 5 -cm-di-
ameter limb, 1.5 m above the ground and slanting
10 degrees from a middle canopy tree. On the
Anjanaharibe-Sud Massif, this diminutive species
was also obtained in both pitfall buckets and el-
evated traps placed at low heights. Voalavo gym-
nocaudus appears to be largely terrestrial but with
scansorial capabilities.

The four individuals taken at 1250 m are adult
males with scrotal testes. The female captured in
the 1625 m zone had large mammae, although not
lactating; three placental scars but no embryos
were noted. A subadult female obtained at 1875
m possessed small mammae and an imperforate
vagina. The mammary formula of Marojejy Voa-
lavo is 1-1-1 (n = 2), as reported for those spec-
imens in the RS d' Anjanaharibe-Sud and mem-
bers of Eliurus.

Comments The Marojejy specimens of Voa-
lavo gymnocaudus possess most of the cardinal
traits advanced for the new genus and species
(Carleton & Goodman, 1998), especially as ob-
served in the absence of a prominent tail tuft, the
small third molars and the unlaminated configu-
ration of M3, the relatively long incisive forami-
na, the reduction of the tegmen tympani, and lack
of an entepicondylar foramen. However, each of
the three cleaned skulls has very thin alisphenoid

struts, a feature not found on the four crania of
the Anjanaharibe-Sud specimens. The molars of
Voalavo are lower crowned than those of Eliurus
species, as suggested by the usual lingual contact
of the posterior lamina on Ml -2, with only little
to moderate wear.

Specimens Examined 11 km NW Mananten-
ina, Antranohofa, 1250 m (FMNH 159725; UA-
SMG 8393); 10.5 km NW Manantenina, 1625 m
(FMNH 159647, 159648); 10.5 km NW Manan-
tenina, 1625 m (FMNH 159726); 11 km NW
Manantenina, at source of Andranomifototra Riv-
er, 1875 m (FMNH 159727).

Discussion

The field survey of small mammals in the PN
de Marojejy, conducted over seven weeks in 1996
at five elevations between 450 and 1875 m, has
generated the first vouchered evidence of native
rodents that inhabit this national park. Seven spe-
cies of endemic Nesomyinae are represented
among the 109 rodent specimens preserved (Table
12-6), including two forms recently described as
new to science from the nearby RS
d' Anjanaharibe-Sud (Voalavo gymnocaudus and
Eliurus grandidieri; Carleton & Goodman, 1998).
Previous faunal reports for the Marojejy Massif
listed only three native species E. cf. myoxinus,
Brachytarsomys albicauda, and Nesomys cf. rufus
(Nicoll & Langrand, 1989; Duckworth & Rako-
tondraparany, 1990). In view of the recent taxo-
nomic changes within Nesomyinae, particularly
the genus Eliurus, it is impossible to confirm ob-
servational field determinations for most members
of the subfamily as reported under the antiquated
nomenclature.

No rodent species new to science was encoun-
tered in the PN de Marojejy. This nondiscovery
repeats that earlier obtained in the RNI
d'Andohahela, where humid forest reaches its
southern limits and where only known forms were
documented using similar trapping equipment and
sampling procedures in 1995 (Goodman, Carle-
ton, & Pidgeon, 1999). In view of the new neso-
myine genera (two) and species (five) described
during the past few years from the eastern forest
(Carleton, 1994; Carleton & Goodman, 1996,
1998), the lack of additional nesomyine variety
from the Andohahela and Marojejy massifs, near-
ly at the polar ends of the island, suggests that
present documentation of rodents occurring in this

CARLETON & GOODMAN: RODENTS

251

biome approximates its actual species diversity.
However, other habitats on the island have yet to
be rigorously surveyed, notably in the west, and
almost certainly other rodent species await dis-
covery.

Trapping Effort and Sampling Confidence

A total of 3,390 trap-nights of sampling effort
were amassed during the Marojejy survey, rang-
ing from 600 to 800 trap-nights per individual ele-
vational zone (Table 12-6). Of the eight species
of rodents captured, all except Rattus rattus are
members of the endemic murid subfamily Neso-
myinae. These animals were found across a broad
cross-section of habitats encompassing over 1400
m of elevation, from lowland forest (450 m), a
zone transitional between lowland and montane
forest (775 m and 1250 m), montane forest (1625
m), and sclerophyllous forest (1875 m). A few
rodents were obtained in pitfall traps installed
mainly to capture lipotyphlans (see Chapter 11),
which were also obtained regularly in live traps
(Table 12-6).

Overall trap success for small mammals (ro-
dents, native and introduced, and insectivores)
varied appreciably among the five elevational
zones. Relatively poor rates (1.4-1.6%) were re-
alized at the lower and highest elevations (450.
775. and 1875 m) and better returns (5.7-7.6%)
at the intermediate sites (1250 and 1625 m) (Table
12-6). Consideration of only Rodentia or only the
first 500 trap-nights per zone discloses the same
elevational profile of trapping success, although
absolute values differ slightly (Tables 12-1 and
12-6). An increase in capture rates for rodents at
middle elevations is proving typical of mountains
surveyed in the eastern humid forests of Mada-
gascar (for example. Goodman & Carleton. 1996.
1998). An exception to this pattern was found on
forested slopes within parcel 1 of RNI
d'Andohahela. where uniformly low levels of trap
success (based on the standardized trapping peri-
od) were obtained for rodents (0.8-3.6% for five
elevations between 440 and 1875 m; Goodman.
Carleton, & Pidgeon, 1999).

The cumulative number of rodent species cap-
tured within each elevational zone generally
reached an asymptote before termination of a
trapping session, usually by the end of the third
trap-night (Fig. 12-4A). At 1250 m. however, the
first example of Eliurus majori was not caught
until the sixth night of trapping, of seven total.
Trap success varied unpredictably from night to
night within each elevational zone. There is nei-
ther evidence of regularly diminishing trap suc-
cess over time at an elevation, at least for the
length of time trapped, nor clear indication that
asymptotes in species accumulation curves are
correlated with declines in capture rates (Fig. 12-
4B).

Notwithstanding the level of effort summarized
above, there is ample reason to question whether
the 1996 survey of the PN de Marojejy was suf-
ficiently comprehensive to document all rodent
species that live there. Individuals of Nesomys, a
comparatively large rodent conspicuous by its di-
urnal activity and reddish fur, were observed dur-
ing the 1996 fieldwork but somehow avoided at-
tempts at capture. Nesomys, as well as Brachytar-
somys, has been previously reported to occur on
the Marojejy Massif (Duckworth & Rakotondra-
parany, 1990). Persuasive evidence for their pres-
ence issues from the RS d'Anjanaharibe-Sud, a
reserve just to the southwest of Marojejy and part
of the same northern highland complex (Fig. 12-
5), where samples of both genera, identified as the
species N. rufus and B. albicauda, were collected
in 1994 (Goodman & Carleton, 1998). Except for
these two, the RS d'Anjanaharibe-Sud otherwise
contains the same species reported here for the PN
de Marojejy, at comparable elevations and in
comparable habitats (see below). Based on this
ecological complementarity and geographic prox-
imity, the aforementioned species of Brachytar-
somys and Nesomys should be expected as inhab-
itants of the PN de Marojejy. The possibility re-
mains that some Nesomys spotted in the reserve
represent the species N. audeberti, a form known
from lowland areas of the eastern humid forest
(see Goodman & Carleton. 1998. p. 214, for dis-
cussion).

Fig. 12-4. Plots of trap-nights against the cumulative number of rodent species (including Rattus rattus) obtained
(A) and against the number of individuals trapped (B) for each of five elevational zones surveyed within the PN de
Marojejy.

252

FIELDIANA: ZOOLOGY

CO
CD

O
CD
Q.
CO

CD
-Q

E

3
C

>

+ i
JO

O

A)

i 1 1 r 1 1 r

100 200 300 400 500 600 700 800 900

Trap-nights

16

CD

2 12

Q.
CO

10

CO
+-

c

CD 8

-o
o

_ 6
o

CD
-Q

E

3

A

*,

. A, f

*,

,'

- a

o -- v

-A"""

-><*'...

"^6

B)

100 200 300 400 500 600 700 800 900

Trap-nights

CARLETON & GOODMAN: RODENTS

253

I I
44 E 46 E

I
48 E

I

50 E

- 12 S

500 m contour

Montagne

mm 1000 -1500 m

XV

v/'d'Ambre

1500 -2000 m

>2000m

<0j-

u.r

- 14 S

Tsaratanana
Marojejy

Anjanaharibe-Sud

\

&

- 16 S

Baie
d'Antongil

V

- 18 S

Andranomay

^

Ankaratra

h 20 s

Fig. 12-5. Northern and central Madagascar, illustrating the close proximity of the PN de Marojejy and RS
d'Anjanaharibe-Sud in the island's northern highlands and other places mentioned in the text.

Elevation and Rodent Associations

Elevational Distribution None of the eight
rodent species found in the PN de Marojejy was
trapped in all five altitudinal zones surveyed (Ta-
ble 12-9). Eliurus grandidieri, E. majori, and
Voalavo gymnocaudus were recorded in the upper
three zones, between 1250 and 1875 m, and these
three species appear to occur across the widest

elevational belt of the native species living on the
eastern slopes of the massif. The few examples of
Voalavo were obtained in live traps only at 1625
m and in pitfall traps at 1250 and 1875 m. Wheth-
er the species occurs in low densities or behaves
so as to avoid live traps is uncertain.

Three nesomyines and the one murine were
found in two contiguous elevational zones: Eli-
urus webbi at 450 and 775 m, E. minor and Ne-

254

FTELDIANA: ZOOLOGY

Table 12-9. Elevational occurrence of rodents in the PN de Marojejy during the 1996 survey based on live traps,
pitfall traps, and sight observations.

Species

450 m

775 m

1250 m

1625 m

1875 m

Murinae

Rattus rattus

+

+

Nesomyinae

Eliurus grandidieri

+

+

+

Eliurus majori

+

+

+

Eliurus minor

+

+

+

Eliurus tanala

+

Eliurus webbi

+

+

Gymnuromys roberti

+

Nesomys cf. rufus

s

s

Voalavo gymnocaudus

+

+

+

Total species

1

4

7

5

3

Total native species

1

4

6

4

3

S = sight record with no associated voucher specimen.

somys cf. rufus at 775 and 1250 m, and Rattus
rattus at 1250 and 1625 m. Eliurus minor is
known to have a wider altitudinal distribution
elsewhere in Madagascar. In the northeastern re-
gion, the species has been recorded near sea level
in the vicinity of Antongil Bay (Carleton, 1994);
near the southern extent of the eastern humid for-
est, in the RNI d'Andohahela, it was captured be-
tween 810 and 1875 m (Goodman, Carleton, &
Pidgeon, 1999). Introduced R. rattus proved to be
exceptionally rare in traps as compared to other
sites recently censused in the eastern humid for-
est. In the nearby site of Anjanaharibe-Sud, for
instance, it was captured in all four elevations
studied between 875 and 1950 m and in both sec-
ondary and pristine forest.

The other two nesomyine species captured in
the PN de Marojejy originated from single zones.
One, Gymnuromys roberti, taken at 1250 m, is
notably difficult to trap and wherever found has
never been recorded in large numbers. At other
sites it has been documented to occur across a
broader elevational range, from the upper reaches
of lowland rain forest into upper montane forest
(e.g., Goodman & Carleton, 1996). The second
species, the usually common Eliurus tanala, was
inexplicably captured only twice and only at 775
m, but it is well known to broadly inhabit low-
land-montane transition and montane communi-
ties at other places in the eastern humid forest
(Goodman & Carleton, 1996, 1998; Goodman,
Carleton, & Pidgeon, 1999). Illuminated by the
now substantial information base on nesomyine
distributions, these examples underscore the need
to temper conclusions on limited elevational oc-

currence, given the stochastic interplay of season-
ally changing population densities, trapping like-
lihood, and the brevity of the field survey (6-8
days at any one elevational level).

No rodent species was recovered in traps
placed in the open grassland above tree line of the
1875 m zone. Of the 100-trap, 540-m-long tran-
sect at this altitude, 49 stations extended through
sclerophyllous forest, 10 across the ecotone be-
tween sclerophyllous forest and grassland, and 4 1
into the nonwooded grassland proper. Voalavo
gymnocaudus, Eliurus grandidieri, and E. majori
were obtained (the first in a pitfall trap) in scler-
ophyllous forest and only E. grandidieri in the
ecotone traps. The cap of alpine habitat occurs on
the Marojejy Massif between about 1850 m and
the summit at 2132 m, and the absence of rodents
wholly restricted to this highland zone conforms
to findings for other tall mountains so far studied
on Madagascar (see Goodman, Carleton, & Pidg-
eon, 1999, and below).

Species Richness The elevational distribution
of rodent species richness in the PN de Marojejy
shows a hump-shaped profile, with fewer species
(N = 1-3) co-occurring at the lowest and highest
zones surveyed and more (N = 4-7) at middle
heights (Table 12-9). The greatest number of na-
tive nesomyines, six species, was recorded at
1250 m, a number and elevation comparable to
biodiversity maxima documented for other moun-
tains surveyed in the eastern humid forest seven
species at 1625 m in the RNI d'Andringitra
(Goodman & Carleton, 1996; Goodman & Raso-
lonandrasana, in press), six species at 1260 m in
the RS d'Anjanaharibe-Sud (Goodman & Carle-

CARLETON & GOODMAN: RODENTS

255

Table 12-10. Estimated biomass (g) of rodents trapped along an elevational transect in the PN de Marojejy.*

Species

450 m

775 m

1250 m

1625 m

1875 m

Rattus rattus
Eliurus grandidieri
Eliurus majori
Eliurus minor
Eliurus tanala
Eliurus webbi
Gymnuromys roberti
Voalavo gymnocaudus

Total biomass (g)

Total biomass (g)
excluding Rattus

Total no. of
species

300

300

300

114

114

1,479

612

306

1,540

110

44

44

206

225

475

475

162

1,799

1,685

22
2,288

2,174

416

416

* Summations from average weight of adults captured over the first 500 trap-nights within each zone.

ton, 1998), and five species at 1200 m in the RNI
d'Andohahela (Goodman, Carleton, & Pidgeon,
1999). The mid-elevational bulge in species rich-
ness within the PN de Marojejy thus mirrors an
elevational pattern common to a variety of organ-
isms on mountains in tropical latitudes (see Good-
man, Carleton, & Pidgeon, 1999, for discussion,
with emphasis on Madagascar and the eastern hu-
mid forest biome).

Not surprisingly, biomass calculations, as based
on animals captured during the first 500 trap-
nights, exhibit the greatest values over the middle
elevations (1250 and 1625 m) and decline mark-
edly toward the lowest and highest elevations
sampled (Table 12-10). At both middle sites, a
single species of Eliurus (E. grandidieri at 1250
m and E. majori at 1625 m) accounted for more
than half of the biomass of captured rodents. The
mid-elevational bulge in standing biomass of ro-
dent populations logically corresponds to both the
number of species and number of individuals
trapped, figures which were greatest at these mid-
dle heights (Tables 12-6 and 12-9). Certain spe-
cies were necessarily excluded from the biomass
comparisons among all elevations where known,
due either to pitfall recovery (Voalavo gymnocau-
dus), visual record (Nesomys rufus), or capture af-
ter the standard trapping period (Eliurus majori at
1250 m). Their absences bias any closer exami-
nation of the relationship between biomass, spe-
cies richness, and population size on Marojejy.

Intra- and Interspecific Differences in Re-
production The adult sex ratio of most rodent
species obtained on Marojejy, as inferred from
trap returns, does not appear to differ substantially

from 1:1 (Table 12-8). For most taxa, however,
sample size is too small to meaningfully test for
significance. A notable exception is E. majori,
nearly two-thirds of the individuals captured be-
ing male. In the RS d'Anjanaharibe-Sud, only E.
grandidieri showed a slightly skewed population
sex ratio, in this instance biased toward females,
based on trap captures. The rodent faunas on the
two massifs are nearly alike in specific composi-
tion, and no pattern in departure from even sex
ratios is detectable for any species common to
both sites. Whatever differences emerged can be
plausibly attributed to the vagaries of trapping
success.

Since a maximum lag of only 10 days separated
the trapping sessions in the five elevational zones,
initiated consecutively from the lowest to highest,
and since age-class recognition does not address
"subadult" physiological fertility, we cannot as-
sess the incidence of breeding with respect to
broader seasonal patterns and elevational trends.
At the time of the survey (October and Novem-
ber), external signs of reproduction were apparent
for all rodent species and across most elevational
zones where sample size is appreciable (Table 12-
8). Moreover, the proportion of subadults was rel-
atively high in the total catch of Eliurus grandi-
dieri and E. webbi, about 38% and 64%, respec-
tively. The relative abundance of immature ani-
mals and the number of actively breeding adults
within each elevational zone where these two spe-
cies were captured suggest that reproduction for
them was ongoing during the period of this sur-
vey. Among examples of E. majori, on the other
hand, few subadults were present, but a majority

256

FIELDIANA: ZOOLOGY

of adults (87%) exhibited some condition imply-
ing active reproduction. In other species, samples
sizes are plainly insufficient to sustain insights on
breeding status of their populations. Although
conducted two years earlier, the survey of the RS
d'Anjanaharibe-Sud (October-November 1994)
occurred during the same calendar period as that
in the PN de Marojejy (October-November
1996). There, most nesomyine species, in partic-
ular E. grandidieri and E. majori, showed high
percentages of subadults relative to adults.

Information on the general natural history of
Madagascar's native rodents has improved mark-
edly in recent years, mainly through coordinated
field surveys in faunistically poorly understood
regions of the eastern humid forest (Goodman &
Carleton, 1996, 1998; Goodman et al., 1997;
Goodman, Carleton, & Pidgeon, 1999). In spite of
this resurgent attention, detailed knowledge about
the reproductive biology of any one species at the
population level has advanced little. The annual
timing and relative brevity of the surveys, most
conducted in the last quarter of the year (October-
December) and over the course of 5-10 days at a
given elevation, prejudice inferences about the
onset of reproduction, its cyclical regularity (or
not), and relationship to environmental rhythms
(rainfall patterns, fruiting cycles, insect fluctua-
tions, etc.). By the criteria adopted (males with
scrotal testes, females with embryos or lactating,
prevalence of subadults), we can say that moder-
ate to high levels of rodent reproduction were oc-
curring in all of these protected areas for the quar-
ter-year period of the surveys. Better understand-
ing of annual nesomyine reproductive patterns
and breeding systems, as well as appreciation of
elevational or altitudinal variation within and
among species, can only be addressed through
site-specific field investigations, undertaken over
a longer period and employing methodologies
such as a grid-based, mark-and-release protocol.

The Rodent Faunas of the PN de Marojejy
and RS d'Anjanaharibe-Sud

Elevational Concordance An initial goal
for surveying small mammals on different moun-
tains throughout Madagascar's eastern humid for-
est, which covers a latitudinal swath of 12 de-
grees, was to assess the relationship between spe-
cies diversity and geophysical gradients like ele-
vation and latitude. Such gradients, and the many
abiotic environmental factors that covary with

them (e.g., rainfall, temperature, insolation, evap-
oration), potentially mold the composition and
productivity of biological communities dwelling
along mountain slopes (see, for example, Rahbek,
1997; Rosenzweig, 1992; Rosenzweig & Abram-
sky, 1993). The validity of these comparisons crit-
ically hinges on appropriate controls that will al-
low biological patterns to be distinguished from
stochastic variation. If the pattern of rodent dis-
tributions on mountains in eastern Madagascar
corresponds in part to gradients associated with
latitude, then elevational affinities of these organ-
isms on nearby mountains at the same latitude
should be expected to closely conform.

The Marojejy and Anjanaharibe-Sud massifs,
separated by a few tens of kilometers and one-
half degree of latitude in the northern highlands
(Fig. 12-5), offer a natural circumstance to ex-
plore this expectation. The rodent surveys of these
two mountains were basically conducted during
the same period: Anjanaharibe-Sud from 15 Oc-
tober to 30 November, 1994, and Marojejy from
4 October to 20 November, 1996. The same line-
transect sampling, trapping methods (live traps
and pitfalls), and baits were used at each reserve.
On both massifs, the surveys were performed
along their eastern-southeastern slopes, across
about 1200 to 1400 m of vertical relief (875-1950
m in Anjanaharibe-Sud; 450-1875 m in Maroje-
jy). The summits of the two mountain systems rise
to nearly equivalent heights, that of Anjanaharibe-
Anivo to 2064 m and Marojejy to 2132 m. The
similar vertical relief of the two mountains lends
further credence to the comparison, since Good-
man, Carleton, & Pidgeon (1999) have disclosed
a strong correlation between elevation at the sum-
mit and that elevation where most rodent species
co-occur; they attributed the relationship to a
combination of orographic variables and their
multifaceted effects.

An exception to their similarity involves the
southeastern side of the RS d'Anjanaharibe-Sud,
whose lowest boundary ends at 875 m, above low-
land rain forest proper like that sampled in the PN
de Marojejy (at 450 m where only Eliurus webbi
was caught). Pair wise comparisons between the
two massifs are therefore restricted to the four
more or less coordinate elevational samples, as fol-
lows (Marojejy and Anjanaharibe-Sud listed re-
spectively): 775 and 875 m, 1250 and 1260 m,
1625 and 1550 m, and 1875 and 1950 m.

The elevational limits and sympatric coinci-
dence of nesomyine species are strongly concor-
dant on the two mountains (Table 12-11). For all

CARLETON & GOODMAN: RODENTS

257

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258

FIELDIANA: ZOOLOGY

Table 12-12. Comparison of rodent species and elevational ranges (in meters) documented to date for comparably
censused regions in eastern humid forests.

Montagne

Anjanaharibe-

d'Ambre 1

Marojejy 2

Sud 3

Andringitra 4

Andohahela 5

Species

340-1350 m

450-1875 m

875-1950 m

720-2450 m

440-1875 m

Mus musculus

2050

Rattus rattus

340-1350

1250-1625

875-1950

810-

-2450

810-1875

Brachytarsomys albicauda

875

Brachyuromys betsileoensis

1990-

-2450

Brachyuromys ramirohitra

1210-

-1990

Eliurus grandidieri

1250-1875

1260-1550

Eliurus majori

1000-1350

1250-1875

1260-1950

1210-

-1990

1200-1500

Eliurus minor

1000

775-1250

875-1260

720-

-1625

810-1875

Eliurus tanala

775

1260

810-

-1625

810-1500

Eliurus webbi

650-1000

450-775

875

720-

-810

440-810

Gymnuromys roberti

1250

1260

720-

-1625

1200

Monticolomys koopmani

1625-

-1990

1875

Nesomys rufus

775-1250

1260-1950

810-

-1990

810-1200

Voalavo gymnocaudus

1250-1875

1950

Number of species

of Nesomyinae

3

8

9

9

7

1 Raxworthy and Nussbaum (1994), Goodman et al. (1996).

2 This study. The listing of Nesomys cf. rufus is based on sight records.

3 Goodman and Carleton (1998). This tabulation does not include collections made by Franco Andreone on the
western slopes of the Anjanaharibe-Sud Massif.

"Goodman and Carleton (1996); Langrand and Goodman (1997); Goodman and Rasolonandrasana (in press).
5 Goodman, Carleton, and Pidgeon (1999).

36 possible pairwise comparisons (nine species by
four elevational zones), no significant contrast in
pattern of specific occurrence can be demonstrat-
ed between the neighboring massifs (Wilcoxon
signed-ranks test, z = -0.38, two-sided P =
0.71). Or, stated in a complementary way, the cor-
relation between a species presence and its high-
est elevation of occurrence on the two mountains
is highly significant (Spearman's p = 0.82, P =
0.011). Further, that elevational belt with greatest
species diversity is the same on Marojejy and An-
janaharibe-Sud (1250-1260 m), a zone corre-
sponding to lower montane forest. Indeed, the co-
presence, or joint absence, of a species within an
elevational zone on the two massifs is so com-
monplace that the lack of complementarity is rea-
sonably suspect as sampling error. The latter in-
stances involve the infrequently captured species
at each reserve, whose numbers seemed to be un-
usually low (Eliurus tanala) or whose habits may
diminish conventional live-trapping success (Voa-
lavo gymnocaudus). The other notable exception,
and the only diurnal species at these two locali-
ties, Nesomys rufus, was commonly trapped from
1260 to 1950 m at Anjanaharibe-Sud, but was ob-
served only at 775 and 1250 m in Marojejy. The
contrast between Anjanaharibe-Sud and Marojejy

in abundance (and catchableness) of Nesomys ru-
fus exposes the weakness of drawing conclusions
from single-point field surveys conducted in dif-
ferent years. Level of reproduction in Nesomys,
and other Nesomyinae, may be sensitive to a host
of other environmental cues, which may vary in
timing and degree between years and which may
not simply track calendrical divisions.

The agreement between the rodent faunas on
Marojejy and Anjanaharibe-Sud in elevational
ranges and botanical associations reinforces the
emerging picture of nesomyine distributions with-
in the eastern humid forest biome (Table 12-12).
Eliurus webbi is a species of lowland forest. At
the upper limits of lowland forest or in transitional
settings, it may occur in sympatry with E. tanala.
Although E. tanala was found only at the upper
limit of lowland forest in the PN de Marojejy (775
m), elsewhere it is known to extend from this belt
into lower to middle montane forest proper. Ele-
vational records of E. minor and Nesomys rufus
typically bridge lowland and montane forest hab-
itats, a distribution that may also characterize
Gymnuromys roberti and Brachytarsomys albi-
cauda across the eastern forest block insofar as
intimated by their very limited trapping records.
Prime montane forest and contiguous areas of

CARLETON & GOODMAN: RODENTS

259

sclerophyllous montane forest harbor E. rnajori,
E. grandidieri, and Voalavo gymnocaudus, at
least in the northern highlands.

Nesomyine species broadly distributed within
eastern humid forest, like Eliurus majori and E.
webbi, demonstrate remarkable elevational fidelity
throughout their ranges, from Anjanaharibe-Sud
and Marojejy in the north to Andringitra and An-
dohahela in the south (Table 12-12). The apparent
lack of notable shifts in elevational occurrence of
these rodent species, across a north-south differ-
ence of nearly 10 degrees, suggests an absence of
latitudinal dampening of vegetational zones on
these mountain slopes. Nevertheless, understand-
ing of variation of Malagasy plant communities
with latitude and altitude still requires much de-
tailed study (see Lewis et al., 1996; Messmer et
al., this volume). Montagne d'Ambre (Fig. 12-5),
with its unique geological history and remote lo-
cation at the extreme northern tip of Madagascar
(Goodman et al., 1996), seems to be an exception
in that the lowland species E. webbi and the mon-
tane species E. majori do co-occur in montane
forest (1000 m) below its low summit (1350 m).

The Alpine Zone Broad expanses of natural
open habitat above tree line exist on Madagascar's
highest mountains, and such alpine vegetation oc-
curs on the uppermost slopes of the Marojejy
Massif and to a much lesser extent on the Anjan-
aharibe-Sud Massif. At about 1800 m, dense
mossy forest on the Marojejy Massif diminishes
in stature and the woody vegetation is dominated
by species with sclerophyllous leaves. Farther up-
slope, about 1850 m, continuous sclerophyllous
forest grades into a patchwork of forest and cri-
coid grassland; the latter habitat is characterized
by species of Graminaceae and low-growing
woody plants, predominantly Ericaceae. Dense
bamboo stands are interdigitated across these eco-
tones. Around 1875 m and above, ericoid grass-
land is common and relatively extensive, except
near the summit (2132 m), where grassland inter-
mingles with areas of exposed rock sparsely cov-
ered with geophytic plants. On the upper slopes
of Anjanaharibe-Anivo, the tree line occurs
around 1980 m, below which is dense mossy for-
est of distinctly sclerophyllous character. Erica-
ceous habitat above tree line is altitudinally less
extensive than on Marojejy but is visibly more
dense and bushy in growth form, with large areas
dominated by Graminaceae and patches of bare
rock near the summit. Botanists believe that the
ericoid grassland on Marojejy is natural and lacks
extensive anthropogenic modifications that char-

acterize some parts of the summital zones of the
Tsaratanana and Andringitra massifs (e.g., Guil-
laumet et al., 1975; Nicoll & Langrand, 1989).

No native rodent was trapped on either of these
northern mountains in open habitats above tree
line. On the upper slopes of Marojejy, Eliurus
grandidieri, E. majori, and Voalavo gymnocaudus
were all captured below or just up to tree line, but
not into the ericoid grassland. The native rodent
fauna below the mossy forest-ericoid bush eco-
tone on Anjanaharibe-Anivo is similar to that of
Marojejy, except that E. grandidieri was not
trapped above 1550 m and Nesomys rufus was
obtained in sclerophyllous forest just below tree
line at 1950 m. Although native rodents are not
known to occupy alpine vegetation on these
northern mountains, birds (see Chapter 10) and
tenrecs (see Chapter 1 1 ) do occur in this nonfor-
ested zone.

The apparent absence of native rodents in the
ericoid zones of Marojejy and Anjanaharibe-Sud
contrasts with the Andringitra Massif, near the
southern end of the Central High Plateau, where
the vole-like genus Brachyuromys is abundant in
similar habitat at high elevations (Goodman &
Rasolonandrasana, in press). This genus is rela-
tively widespread across upland locales on the
Central High Plateau: one species, B. ramirohitra,
appears to be restricted to the southern portion of
the central highlands (900-1625 m), and the sec-
ond, B. betsileoensis, is more widely distributed,
from Andringitra in the south to the vicinity of
Lac Alaotra in the north (900-2450 m). This latter
species is known to occur in human-modified hab-
itats. The Andringitra Massif today bears ericoid
grassland that is closest to those mountains sur-
rounding the Andapa Basin, some 800 km to the
north, although the Ankaratra Massif (Fig. 12-5)
plausibly supported such vegetation prior to hu-
man disturbance in the form of forest clearance
and fires. Brachyuromys betsileoensis has also
been obtained in grassy wet meadows on Mount
Ankaratra at elevations between 1705 and 1980
m (Carleton & Schmidt, 1990), a zone that prob-
ably bridged the upper limit of montane forest and
open alpine vegetation before massive destruction
of the natural habitats on this mountain. To date,
Brachyuromys has not been found in the northern
(Goodman & Carleton, 1998; this study) or south-
ern highland blocks (Goodman, Carleton, & Pidg-
eon, 1999) that stand apart from the Central High
Plateau. However, its possible occurrence in the
broad expanse of ericoid grassland covering the

260

FIELDIANA: ZOOLOGY

summital area of the Tsaratanana Massif in the
north invites exploration.

Literature Cited

Burney, D. A. 1987. Pre-settlement vegetation changes
at Lake Tritrivakely, Madagascar. Palaeoecology of
Africa, 18: 357-381.

1997. Theories and facts regarding Holocene

faunal inventory of the eastern slopes of the Reserve
Naturelle Integrale d'Andringitra, Madagascar: With
reference to elevational variation. Fieldiana: Zoology,
85: 257-283.

1998. The rodents of the Reserve Speciale

environmental change on Madagascar before and after
human colonization, pp. 75-89. In Goodman, S. M.,
and B. D. Patterson, eds., Natural change and human
impact in Madagascar. Smithsonian Institution Press,
Washington, D.C.

Carleton, M. D. 1994. Systematic studies of Madagas-
car's endemic rodents (Muroidea: Nesomyinae): Re-
vision of the genus Eliurus. American Museum Nov-
itates, 3087: 1-55.

Carleton, M. D., and S. M. Goodman. 1996. System-
atic studies of Madagascar's endemic rodents (Muro-
idea: Nesomyinae): A new genus and species from the
Central Highlands. In Goodman, S. M., ed., A floral
and faunal inventory of the eastern slopes of the Re-
serve Naturelle Integrale d'Andringitra, Madagascar:
With reference to elevational variation. Fieldiana: Zo-
ology, n.s., 85: 231-256.

. 1998. New taxa of nesomyine rodents (Muro-
idea: Muridae) from Madagascar's northern highlands,
with taxonomic comments on previously described
forms. In Goodman, S. M., ed., A floral and faunal
inventory of the Reserve Speciale d'Anjanaharibe-
Sud, Madagascar: With reference to elevational vari-
ation. Fieldiana: Zoology, n.s., 90: 163-200.

Carleton, M. D., and D. F. Schmidt. 1990. Systematic
studies of Madagascar's endemic rodents (Muroidea:
Nesomyinae): An annotated gazetteer of collecting lo-
calities of known forms. American Museum Novita-
tes, 2987: 1-36.

Duckworth, J. W, and F. Rakotondraparany. 1990.
The mammals of Marojejy, pp. 54-60. In Safford, R.,
and Duckworth, J. W, eds., A wildlife survey of the
Marojejy Nature Reserve, Madagascar. International
Council for Bird Preservation, Cambridge.

Ellerman, J. R. 1949. The families and genera of living
rodents, vol. 3, appendix II [Notes on the rodents from
Madagascar in the British Museum, and on a collec-
tion from the island obtained by Mr. C. S. Webb].
British Museum (Natural History), London, v + 210
pp.

Gasse, F, E. Cortijo, J.-R. Disnar, L. Ferry, E. Gil-
bert, C. Kissel, F. Laggoun-Defarge, E. Lallier-
Verges, J.-C. Miskovsky, B. Ratsimbazafy, F. Ran-

AIVO, L. ROBISON, P. TUCHOLKA, J.-L. SAOS, A. SlFFED-

ine, M. Taieb, E. Van Campo, and D. Williamson.
1994. A 36 ka environmental record in the southern
tropics: Lake Tritrivakely (Madagascar). Comptes
Rendus de l'Academie des Sciences, serie II, 318:
1513-1519.
Goodman, S. M., and M. D. Carleton. 1996. The ro-
dents of the Reserve Naturelle Integrale d'Andringitra,
Madagascar. In Goodman, S. M., ed., A floral and

d'Anjanaharibe-Sud, Madagascar. In Goodman, S. M.,
ed., A floral and faunal inventory of the Reserve Spe-
ciale d'Anjanaharibe-Sud, Madagascar: With refer-
ence to elevational variation. Fieldiana: Zoology, n.s.,
90: 201-221.

Goodman, S. M., and D. Rakotondravony. 2000. The
effects of forest fragmentation and isolation on insec-
tivorous small mammals (Lipotyphla) on the Central
High Plateau of Madagascar. Journal of Zoology, Lon-
don, 250: 193-200.

Goodman, S. M., and B. P. N. Rasolonandrasana. In
press. Elevational zonation of birds, insectivores, ro-
dents, and primates on the slopes of the Andringitra
Massif, Madagascar. Journal of Natural History.

Goodman, S. M., M. D. Carleton, and M. Pidgeon.
1999. The rodents of the Reserve Naturelle Integrale
d'Andohahela, Madagascar. In Goodman, S. M., ed.,
A floral and faunal inventory of the Reserve Naturelle
Integrale d'Andohahela, Madagascar: With reference
to elevational variation. Fieldiana: Zoology, n.s., 94:
217-249.

Goodman, S. M., P. D. Jenkins, and M. Pidgeon. 1999.
The Lipotyphla (Tenrecidae and Soricidae) of the Re-
serve Naturelle Integrale d'Andohahela, Madagascar.
In Goodman, S. M., ed., A floral and faunal inventory
of the Reserve Naturelle Integrale d'Andohahela,
Madagascar: With reference to elevational variation.
Fieldiana: Zoology, n.s., 94: 187-216.

Goodman, S. M., A. Andrianarimisa, L. E. Olson, and
V. Soarimalala. 1996. Patterns of elevational distri-
bution of birds and small mammals in the humid for-
est of Montagne d'Ambre Madagascar. Ecotropica, 2:
87-98.

Goodman, S. M., J. Ganzhorn, L. E. Olson, M. Pidg-
eon, and V. Soarimalala. 1997. Annual variation in
species diversity and relative density of rodents and
insectivores in the Pare National de la Montagne
d'Ambre, Madagascar. Ecotropica, 3: 109-118.

Goodman, S. M., D. Rakotondravony, L. E. Olson, E.
Razafimahatratra, and V. Soarimalala. 1998. Les
insectivores et les rongeurs, pp. 80-93. In Rakoton-
dravony, D., and S. M. Goodman, eds., Inventaire
biologique, Foret d'Andranomay, Anjozorobe. World
Wide Fund for Nature Centre d' Information et de
Documentation Scientifique et Technique, Antanana-
rivo, Recherches pour le Developpement, Serie Sci-
ences Biologiques, no. 13.

Green, G. M., and R. W Sussman. 1990. Deforestation
history of the eastern rain forests of Madagascar from
satellite images. Science, 248: 213-215.

Griveaud, P. 1960. Une mission entomologique au Ma-
rojejy. Naturaliste Malgache, 12: 43-55.

Guillaumet, J.-L., J.-M. Betsch, C. Blanc, P. Morat,
and A. Peyrieras. 1975. Etude des 6cosystemes mon-
tagnards dans la region malgache. III. Le Marojezy.
IV. LTtremo, et l'lbity. Geomorphologie, climatologie,
fauna et flore (Campagne RCP 225, 1972-1973). Bul-
letin du Museum National d'Histoire Naturelle, 3rd
series, no. 309, Ecologie Generate, 24: 29-67.

CARLETON & GOODMAN: RODENTS

261

Hawkins, A. F. A., J.-M. Thiollay, & S. M. Goodman.
1998. The birds of the Reserve Spe"ciale
d'Anjanaharibe-Sud, Madagascar. In Goodman, S. M.,
ed., A floral and faunal inventory of the Reserve Spe-
ciale d'Anjanaharibe-Sud, Madagascar: With refer-
ence to elevational variation. Fieldiana: Zoology, n.s.,
90: 93-127.

Humbert, H., and G. Cours Darne. 1965. Carte inter-
nationale du tapis vegetal et des conditions ecolo-
giques. Travaux de la Section Scientifique et Tech-
nique de lTnstitut Francais de Pondichery (hors serie).

Jansa, S. A. 1998. Molecular phylogeny and biogeog-
raphy of Madagascar's native rodents (Muridae: Ne-
somyinae). Ph. D. thesis. University of Michigan, 200
pp.

Jenkins, P. D. 1987. Catalogue of primates in the British
Museum (Natural History) and elsewhere in the Brit-
ish Isles. Part IV. Suborder Strepsirrhini, including the
subfossil Madagascan lemurs and family Tarsiidae.
British Museum (Natural History), London, x + 189
pp.

Langrand, O., and S. M. Goodman. 1997. Inventaire
des oiseaux et des micro-mammiferes des zones som-
mitales de la Reserve Naturelle Integrale
d'Andringitra. Akon'ny Ala, 20: 39-54.

Lewis, B. A., P. B. Phillipson, M. Andrianarisata, G.
Rahajasoa, P. J. Rakotomalaza, M. Randriambol-
olona, and J. F. McDonagh. 1994. A study of the
botanical structure, composition, and diversity of the
eastern slopes of the Reserve Naturelle Integrale
d'Andringitra, Madagascar. In Goodman, S. M., ed.,
A floral and faunal inventory of the eastern slopes of
the Reserve Naturelle Integrale d'Andringitra, Mada-
gascar: With reference to elevational variation. Fiel-
diana: Zoology, n.s., 85: 24-75.

Nicoll, M. E., and O. Langrand. 1989. Madagascar:
Revue de la conservation et des Aires Protegees.
World Wide Fund for Nature, Gland, Switzerland, xvii
+ 374 pp.

Rahbek, C. 1997. The relationship among area, eleva-
tion, and regional species richness in Neotropical
birds. The American Naturalist, 149: 875-902.

Rakotondravony, D., S. M. Goodman, J.-M. Duplan-
tier, and V. Soarimalala. 1998. Les petits mammi-
feres, pp. 197-212. In: Ratsirarson, J., and S. M.
Goodman, eds., Inventaire biologique de la Foret Lit-
torale de Tampolo. Centre d'lnformation et de Docu-
mentation Scientifique et Technique, Antananarivo,
Recherches pour le Developpement, Serie Sciences
Biologiques, no. 14.

Rand, A. L. 1932. Mission Franco- Anglo-Americaine a
Madagascar: Notes de voyage. Oiseau et Revue Fran-
chise d'Ornithologie, 2: 227-282.

. 1936. The distribution and the habits of Mad-
agascar birds: A summary of the field notes of the
Mission Zoologique Franco-Anglo-Americaine a
Madagascar. Bulletin of the American Museum of
Natural History, 72: 143-499.

Raxworthy, C. J., and R. A. Nussbaum. 1994. A rain-
forest survey of amphibians, reptiles and small mam-
mals at Montagne d'Ambre, Madagascar. Biological
Conservation, 69: 65-73.

. 1996. Montane amphibian and reptile commu-

nities in Madagascar. Conservation Biology, 10: 750-
756.

Rosenzweig, M. L. 1992. Species diversity gradients:
We know more or less what we thought. Journal of
Mammalogy, 73: 715-730.

Rosenzweig, M. L., and Z. Abramsky. 1993. How are
diversity and productivity related? pp. 52-65. In Rick-
lefs, R. E., and D. Schluter, eds.. Species diversity in
ecological communities. University of Chicago Press,
Chicago, 414 pp.

Stephenson, P. J., H. Randriamahazo, N. Rakotoari-
son, and P. A. Racey. 1994. Conservation of mam-
malian species diversity in Ambohitantely Special Re-
serve, Madagascar. Biological Conservation, 69: 213
218.

Thomas, O. 1895. On a new species of Eliurus. Annals
and Magazine of Natural History, series 6, 16: 164-
165.

. 1908. A new species of the Mascarene genus

Eliurus. Annals and Magazine of Natural History, se-
ries 8, 2: 453-454.

Voss, R. S., and M. D. Carleton. 1993. A new genus
for Hesperomys molitor Winge and Holochilus mag-
nus Hershkovitz (Mammalia, Muridae) with an anal-
ysis of its phylogenetic relationships. American Mu-
seum Novitates, 3085: 1-39.

Appendix 12-1

Listed below are vouchered specimens of Eli-
urus collected outside of PN de Marojejy that
were used in the taxonomic comparisons and mor-
phometric analyses. They are contained in the fol-
lowing museums: British Museum (Natural His-
tory), London (BMNH); Field Museum of Natural
History, Chicago (FMNH); Merseyside County
Museums, Liverpool (LMCM); Museum of Com-
parative Zoology, Harvard University, Cambridge
(MCZ); Museum National d'Histoire Naturelle,
Paris (MNHN); National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C.
(USNM); Universitets Zoologisk Museum, Co-
penhagen (UZMC); and Zoologisches Forschung-
sinstitut und Museum Alexander Koenig, Bonn
(ZFMK).

Eliurus majori Antsiranana Province: PN de
la Montagne d'Ambre, 1000-1350 m (AMNH
100687, 100854; FMNH 154535-154539,
154603-154616, 156341-156345); RS

d'Anjanaharibe-Sud, 1260-1950 m (FMNH
154052-154054, 154241-154245, 154266,
154289). Fianarantsoa Province: Ambohimitombo
(BMNH 97.9.1.147; MCZ 45929); RNI
d'Andringitra, 1210-1625 m (FMNH 151661-
151667, 151730-151732, 151752, 151847,

262

FIELDIANA: ZOOLOGY

151848, 151852, 151853, 151855); Anjavidilava
(MNHN 1972.602); RS d'lvohibe, 1575 m
(FMNH 161891, 162073-162075). Toliara Prov-
ince: RNI d'Andohahela, 1200-1500 m (FMNH
156615-156617, 156658).

Eliurus penicillatus Fianarantsoa Province:
Ampitambe (BMNH 97.9.1.148-97.9.1.150,
97.9.1.152, 1939.1892; FMNH 18822; LMCM
A 19.4.98.26; MCZ 45932; MNHN 1897.536,
1909.191; USNM 49672; UZMC 1219, 1224,
7941; ZFMK 82.217, 83.56).

Eliurus webbi Antsiranana Province: PN de la
Montagne d'Ambre, 650-1000 m (MNHN
1957.781, 1970.248, 1970.249, 1973.517,
1973.518; UA-SMG 6755); RS d'Anjanaharibe-
Sud, 875 m (FMNH 154035-154038, 154251,
154252). Toamasina Province: 3 mi E Andrano-
fotsy, 12 km NE Maroantsetra, near sea level
(BMNH 47.1585, 47.1586, 47.1588); Antalaha
(MNHN 1961.179); Antalavia, E side Antongil
Bay, near sea level (BMNH 47.1590); 40 km SW

Maroantsetra, 10 km WSW Maintimbato, 450-
625 m (BMNH 1987.108); 40 km NE Maroantse-
tra, Ankovana, 900 ft (BMNH 47.1589); 8 mi W
Rantabe, Antongil Bay, 500 ft (BMNH 47.1593).
Fianarantsoa Province: 2 km NE Andrambovato,
575-625 m (USNM 449266-449268); 1 km NW
Andrambovato, 875 m (USNM 449269); 12 km
E Ifanadiana (MNHN 1961.216-1961.219); 0.5
km N Kianjavato, 300 m (USNM 448991-
448995); 9 km ESE Kianjavato, 250-500 m
(USNM 449257-449265); RNI d'Andringitra,
720-810 m (FMNH 151680-151686, 151739-
151742, 151884, 151885, 151888, 151891-
151895, 156513, 156523-156526, 156529,
156644); RS d'lvohibe, 900 m (BMNH 47.1594-
47.1599; FMNH 162091); 20 km W Vondrozo
(MNHN 1932.3517, 1957.782); 20 mi S Farafan-
gana, 4 mi from sea (AMNH 119707; BMNH
47.1574-47.1579, 47.1582-47.1584). Toliara
Province: RNI d'Andohahela, 440-810 m
(FMNH 156513, 156523-156526, 156529,
156644).

CARLETON & GOODMAN: RODENTS

263

Chapter 13

Rapid Census of Lemur Populations in the Pare
National de Marojejy, Madagascar

Eleanor Sterling 1 and Karen McFadden 2

Abstract

This chapter discusses the results of a primate survey conducted between 4 October and 20
November 1996 in the Pare National de Marojejy, northeastern Madagascar. Research sites
were established at five elevations 450, 775, 1250, 1625, and 1875 m; surveys were confined
to areas 75 m higher or lower in altitude at each elevation. Species richness and an index of
density were calculated for lemur species, at all sites, by means of both the line transect method
and an experimental density estimation method. Ten lemur species were recorded in the reserve.
Species richness did not differ greatly between the four lower elevational zones but dropped
sharply at the highest elevation. Key taxa found within the reserve include Daubentonia mad-
agascariensis, Propithecus diadema candidus and possibly Phaner furcifer. Other taxa ob-
served in the reserve include Eulemur fulvus albifrons, E. rubriventer, Hapalemur griseus
griseus, Avahi laniger, Cheirogaleus major, Microcebus rufus, and Lepilemur mustelinus.

Resume

Cet article commente les resultats de l'inventaire des primates, conduit entre le 4 octobre et
le 20 novembre 1996, dans le Pare National de Marojejy, dans le Nord-Est de Madagascar.
Les sites de recherche ont ete repartis sur cinq niveaux d'altitude: 450, 775, 1250, 1625, et
1875 m, l'inventaire etant limite a l'interieur d'une bande de 75 m en dessus et au-dessous de
chaque niveau d'altitude. La richesse specifique et l'indice de la densite des lemuriens ont ete
mesures dans tous les sites par 1' utilisation simultanee de la methode des transects et d'une
methode experimentale pour estimer la densite. Dix especes de lemuriens ont ete enregistrees
dans le pare. La richesse specifique ne differe pas beaucoup entre les quatre plus basses zones
d' altitudes, mais elle baisse considerablement au niveau de 1' altitude la plus elevee. Les groupes
taxinomiques cles trouves dans le pare comprennent Daubentonia madagascariensis, Propithe-
cus diadema candidus et peut etre Phaner furcifer. Les autres especes observees dans le pare
incluent Eulemur fulvus albifrons, Eulemur rubriventer, Hapalemur griseus griseus, Avahi lan-
iger, Cheirogaleus major, Microcebus rufus, et Lepilemur mustelinus.

Introduction

1 Center for Biodiversity and Conservation, American Madagascar's northeastern forests, representing

Museum of Natural History, New York, NY 10024, a ma j ority of the COU ntry's remaining intact rain-

U '^Department of Anthropology, Yale University, New f rest tracts > harbor hi g h levels of floral and fau '

Haven, CT 06511, U.S.A. nal diversity and endemism. Although the lemur

STERLING AND MCFADDEN: LEMURS 265

populations are a prominent feature of these rain-
forest areas, comprehensive survey work in these
communities remains to be done. Meanwhile,
conversion of forest tracts for agricultural use
continues to fragment the habitats of many spe-
cies, and hunting pressures directly affect the pri-
mate populations.

The Pare National (PN) de Marojejy is com-
posed primarily of moist montane and sclero-
phyllus montane forest. This area serves as a wa-
tershed for numerous outlying areas. Accurate
baseline inventory data are critical for establish-
ing an effective management plan for this re-
serve's biota. Ideally, such information would
come from intensive, long-term surveys. Howev-
er, hunting and resource extraction pressure on the
reserve, coupled with inadequate protection mea-
sures and limited resources, necessitate that more-
rapid and efficient methods of assessment be used.

As part of an extensive program to assess bio-
diversity in forested areas, a multitaxa survey and
inventory expedition was undertaken in the PN de
Marojejy from early October to mid-November
1996. Data on primate species richness and abun-
dance were collected in the southeastern sector of
the reserve. Earlier surveys identified a total of
nine species of primates in the PN de Marojejy
(Humbert, 1955; Guillaumet et al., 1975; Benson
et al., 1976, 1977; Tattersall, 1982; Nicoll & Lan-
grand, 1989; Safford & Duckworth, 1990), includ-
ing endangered species such as the Sifaka (Propi-
thecus diadema) and the Aye-aye (Daubentonia
madagascariensis). Sightings from the Reserve
Speciale (RS) d'Anjanaharibe-Sud, 20 km to Ma-
rojejy *s southwest (Schmid & Smolker, 1998), and
the PN de Masoala, approximately 100 km to the
southeast (Sterling & Rakotoarison, 1998), suggest
that the PN de Marojejy could possibly include
other endangered primate species, such as the
Hairy-eared Dwarf Lemur (Allocebus trichotis). In-
formation about the distribution and density of pri-
mate species in the PN de Marojejy and the degree
of threat to these populations represents important
data for the Association National pour la Gestion
des Aires Protegees, which manages the reserve in
the context of an integrated conservation and de-
velopment project, under the direction of the World
Wide Fund for Nature.

Methods

Study Area

The PN de Marojejy undergoes two extensive
wet seasons the intense November to April sea-

son, peaking in January, when cyclones occur, and
the lesser June to August season, which is the
time of year with the lowest temperatures in other
northeastern rainforests (Sterling. 1993). In the
present study, survey zones were established at
five sites along an elevational gradient. Each ele-
vational zone (450. 775. 1250. 1625. and 1875 m)
encompassed an area 75 m in altitude, to create
nonoverlapping 150 m bands.

Census Methods

Surveys were conducted between 4 October
and 20 November 1996. During the 10 days we
spent at each elevational zone, two survey meth-
ods were used, including the line transect method
and an experimental technique that used both
transect and nontransect observations. Previous
expeditions of similar length had indicated that
the line transect method may not yield enough
data in 10 days to allow for the calculation of
density estimates (Sterling & Rakotoarison.
1998); consequently, a second method was tested
in this study.

The line transect method was used along four
to six different trails at each elevational zone.
Some of the trails were pre-existing; others were
opened and marked for this study. Sightings were
made by one observer while walking these trails
for distances varying from 125 to 1475 m. Diur-
nal surveys were conducted between 500 and
1100 and between 1400 and 1700 hr, for an av-
erage of 6 hours per day. Nocturnal surveys oc-
curred between 1 830 hr and dawn, for an average
of 4 hours on any one night. Transects were
walked at a steady speed of 1 km per hour during
the day and 0.75 km per hour at night. Low-power
headlamps were used on night census walks to
detect nocturnal species, whose eyeshine reflects
the light. Brighter lights were then used to illu-
minate the animal for identification. Weather per-
mitting, each trail was censused at least three
times (ranging from two to six times per day/
night) during the 10 days, for both diurnal and
nocturnal censuses, but never consecutively dur-
ing the same day or night session. When groups
were detected along the transect trail, the perpen-
dicular distance from the animal to the transect
was estimated for the first individual in each
sighted group. The distance and the angle from
the observer to the animal were also recorded, to
test the accuracy of our perpendicular distance es-
timates.

266

FIELDIANA: ZOOLOGY

Table 13-1. Species richness by elevational zone for transect and nontransect data, October-November 1996, PN
de Marojejy.

Presence at elevational

zone i m )

Species

450

775

1250

1625

1875

Eulemur fulvus albifrons

X

X

X

X

Eulemur rubriventer

X*

X

X

X

Hapalemur griseus griseus

X

X

X

X

Propithecus diadema candidus

X

X

X

Avahi laniger

X

X

X

Cheirogaleus major

X

X

X

X

Microcebus rufus

X

X

X

X

X

Lepilemur mustelinus

X?*

X

X

X

Phaner furcifer

X?*

Daubentonia madagascariensis

X

X

X

X

X

Total no. of species

8

8

9

8

3

Species heard but not seen at this elevation.

When animals were sighted, pelage color, body
size, height in tree, type of habitat, location along
the transect, and behavioral and feeding infor-
mation were noted, for each individual, as often
as possible. For lemurs detected in groups (mainly
diurnal species) the total number and demograph-
ic composition of the group was recorded.

The second census method (a modification of
Merenlender et al., 1998) incorporated both tran-
sect and nontransect sightings of animals, to ar-
rive at a density estimate. All sightings of lemurs
by primatologists and other experienced primate
observers on the expedition, whether on- or off-
trail, were recorded, and morphological and be-
havioral information similar to that collected on
the transects was noted. Distance and angle were
taken from the nearest topographic feature if there
was no nearby trail. Taxa seen more than three
times were plotted on a map of the transect area.
The outermost of these points were plotted by
means of a global positioning system (GPS) and
were corrected via topographical, hydrological,
and altimeter data. Area calculations for density
estimates were created by calculation of the min-
imum convex polygon linking up the outer points.

Groups were identified by means of one of two
methods. First, several groups contained individ-
uals with distinctive pelage characters. This in-
formation was combined with information on
number of individuals and sex ratio to identify
groups. Second, data on home range area for pop-
ulations of the same species in other eastern rain-
forest reserves (Mittermeier et al., 1994) were
used to estimate which of the sightings were with-
in the species' average home range and which
were outside the range. Sightings farther apart

than the average home range size for the species
were presumed to be distinct animals or groups
and were used to calculate a maximum population
size. Sightings within home range size were treat-
ed as the same group or animal and were used to
calculate a minimum population size.

During all observation periods, we looked for
secondary signs of lemur presence, including
nests, fallen fruit with tooth comb or incisor
marks, and Aye-aye feeding holes in dead trees,
in bamboo, or in Canarium (Burseraceae) seeds.

Results and Species Accounts

Ten species were recorded within the reserve,
across all elevations (Table 13-1). Nine of those
species were observed directly, whereas one (pos-
sibly Phaner furcifer) was only heard. None of
the five elevational zones contained all ten spe-
cies. At 450, 775, and 1250 m, the total number
of species remained relatively similar (range, 8-
9), but species composition varied. At the lowest
elevational zone, we did not encounter Lepilemur,
but we heard calls that were either Phaner furcifer
or Lepilemur. At the second camp, we did see
Lepilemur. At the highest elevational zone, only
three lemur species were recorded.

Since the number of sightings at all elevational
zones was insufficient to calculate transect width,
no estimates of lemur densities were made for the
line transect data. Data from line transects are in-
stead presented as number of sightings of lemurs
per kilometer transect (Tables 13-2 and 13-3).
Data for the experimental method are presented

STERLING AND MCFADDEN: LEMURS

267

Table 13-2. No. of individuals seen per kilometer transect for diurnal primate species, PN de Marojejy, October-
November 1996.

Total

distance

walked

(m)

No.

of individual

s of species

No. of

Altitude and site

Efa

Er

Hgg

Pdc

species

450 m

la

2975

1

1

2

lb

2400

5

1

2

lc

3050

0.3

1

2

Id

3200

3

1

Detection distance

6.0

4.6

7.5 1.2

775 m

2a

1000

2b

1600

3

1

2c

5000

1

2

2

2d

2550

2

1

2e

1800

3

1

2

Detection distance

14.5

7.6

5.8

-i-

6.0

12.9 7.9

1250 m

3a

825

4

4

2

3b

1200

4

1

3c

5375

1

1

2

3d

3800

1

1

1

3

Detection distance

5*

14.2

4.0

15*

19.0

6.6

1625 m

4a

2000

1

1

4b

2325

2

2

2

4c

8325

2

1

4d

125

8

1

4e

500

Detection distance

13.3

10.1

10*

13.5

2.1

1875 m

6

5a

875

4

1

5b

1150

1

5c

500

5d

1300

5e

250

6.0

5.7

Detection distance

9

10

11

7

N

Mean group size

3.1

1.9

2.2

!

0.8

3.2 1.7

3.9

1.7

Note Efa = Eulemur fulvus albifrons, Er = E. rubriventer, Hgg =
Propithecus diadema candidus. Detection distance is the mean distance (:
trail, at which lemurs were seen.

* Single sighting at this elevation.

Hapalemur griseus griseus, and Pdc =
standard deviation), perpendicular to the

as maximum and minimum density estimates (Ta-
ble 13-4).

Five lemur traps, consisting of forest clearings
of 20 m in width, with a branch traversing the
open area covered with a spiral trap, were found
at the first elevational zone. In one of these traps,
we recovered the skull of an individual belonging
to Eulemur fulvus. Many species, notably Hapa-
lemur griseus, Propithecus diadema candidus, E.
fulvus albifrons, and E. rubriventer, reacted

strongly to the presence of humans, generally flee-
ing or hiding and alarm calling.

Avahi laniger

Avahi laniger was found at the first three ele-
vational zones and was not observed at 1625 or
1875 m. These lemurs were seen in a pair once.

268

FIELDIANA: ZOOLOGY

Table 13-3. No. of individuals seen per kilometer transect for nocturnal primate species, PN de Marojejy, Oc-
tober-November 1996.

Total

distance
walked

(m)

No. of individuals of species

No. of

Altitude and site

Al

Lm

Mr

Cm

species

450 m

la

2425

0.4

1

lb

1200

1

1

lc

2050

0.5

1

Id

2100

1

1

Detection distance

3.9 1.4

2.5*

Total observations

4

1

775 m

2a

350

3

3

2

2b

800

2c

1475

2d

1600

1

1

2e

900

1

2

2

1

4

2f

150

7

7

2

Detection distance

20*

4.2 4.7

8.8 1.5

6.3 5.5

Total observations

1

3

3

3

1250 m

3a

500

3b

1200

1

2

2

3c

5900

0.2

1

0.2

3

3d

2550

0.4

1

2

Detection distance

11.0 3.6

18*

10.5 3.5

Total observations

3

1

4

2

1625 m

4a

150

7

1

4b

2600

1

4c

375

3

1

4d

375

Detection distance

15.3 8.7

25*

Total observations

3

1

1875 m

5a

825

1

1

5b

275

Detection distance

10*

Total observation

1

Note Al = Avahi laniger, Lm = Lepilemur mustelinus, Mr = Microcebus rufus, and Cm = Cheirogaleus major.
Detection distance is the mean distance ( standard deviation), perpendicular to the trail, at which lemurs were seen.
* Single sighting at this elevation.

but otherwise singly. No young were observed
during this study.

different individuals in the reserve, and no young
were observed during this study.

Cheirogaleus major

Cheirogaleus major was found at the four low-
est elevational zones but was never particularly
common. All individuals were observed singly.
No significant size difference was noted between

Daubentonia madagascariensis

Proximate evidence of Daubentonia madagas-
cariensis was found in all elevational zones. This
species was observed directly only once, during a
nocturnal transect in the 450 m zone. It was seen

STERLING AND MCFADDEN: LEMURS

269

Table 13-4. Absolute density estimates in individuals per square kilometer for minimum convex polygon areas,
October-November 1996, PN de Marojejy.

Individuals per square kilometer, at altitude (m)

450

775

1250

1625

1875

Species

Eulemur fulvus albifrons
Eulemur rubriventer
Hapalemur griseus griseus
Propithecus diadema candidus
Avahi laniger
Lepilemur mustelinus
Microcebus rufus
Cheirogaleus major
Daubentonia madagascariensis

Min. Max. Min. Max. Min. Max. Min. Max. Min. Max.

24

24

37

57

10

20

37

33

47

57

82

*

40

*

10

16

20

16

20

20

7

13

12

*

5

90

10 15

Note. Min. = minimum; Max. = maximum.

* Taxa observed fewer than three times.

t For this species, presence was inferred at these elevations, on the basis of distinctive signs.

for only an instant as it crossed the path in front
of the observers, at ground level, and then dis-
appeared into a bamboo thicket. In the lowest ele-
vational zone, we noted an old Aye-aye nest, and
we saw several stands of bamboo with character-
istics signs of Aye-aye feeding (Duckworth,
1993). In the 775 and 1250 m zones, we noted
remains of Canarium spp. nuts with clear Aye-
aye feeding marks (Iwano & Iwakawa, 1988;
Goodman & Sterling, 1996). In the two upper-
most elevational zones, observers located remains
of deadwood and also of bamboo where Aye-ayes
had fed.

(Asio madagascariensis) pellet collected at the
1625 m site contained the remains of an M. rufus
(S. Goodman, pers. comm.). Given the small
home range size of this owl, the mouse lemur was
mostly likely taken near where the pellet was de-
posited (S. Goodman, pers. comm.). Observed
pelage characteristics: upper parts and dorsum
reddish-brown, thin dorsal black stripe, and dark
brown tail.

Diurnal Censuses

Lepilemur mustelinus

Lepilemur mustelinus was found in the three
middle elevational zones (Table 13-1). Animals
were generally seen singly, although one obser-
vation included two animals, a female and a
young animal, together. Observed pelage charac-
teristics: upper parts mainly chestnut brown, tail
black, and whitish whiskers on some individuals.

Microcebus rufus

Microcebus rufus occurs at all elevational
zones. We often heard but had few direct obser-
vations of this species. Researchers located a nest
just outside the 775 m zone in October 1996. The
nest contained one young mouse lemur with new-
ly opened eyes. At 1250 m, we recorded Micro-
cebus feeding on fruit from a Dypsis sp. An owl

Hapalemur griseus griseus

Hapalemur griseus griseus groups were found
in all but the highest elevational zone. Average
group size in the reserve was 3.2 1.7 individ-
uals. Researchers observed them feeding on a fun-
gus (undetermined) and on wild bananas (Musa-
ceae) in the 450 m zone. Cryptoprocta scat col-
lected in the 1250 m zone and subsequently ana-
lyzed by Steve Goodman contained bones of H.
griseus. Notable pelage characteristics: dark ru-
fous ventrum, and a pronounced white triangle
around nose.

Eulemur fulvus albifrons

Groups of this species were observed in the
first four elevational zones. Average group size in
the reserve was estimated at 3.1 1.9 individuals.
This is low compared with populations in other

270

FTELDIANA: ZOOLOGY

areas, and it may be a sampling artifact due to
lemurs fleeing from humans in the area. At 450
m, researchers observed individuals feeding on
Meleastrum sp., whereas in the 1250 m zone they
were observed eating Cryptocarya sp. the major
fruiting species in this zone other than Canarium,
during the transect study. Young individuals were
seen in 30% of the observed groups. Observed
pelage characteristics: upper and dorsal pelage
color in males varied from dark brown to light
gray.

Eulemur rubriventer

No group of this species was found in the high-
est elevational zone, although groups were ob-
served in all others. One group was recorded on
audiotape in the 450 m zone, but none were ac-
tually seen in this zone. Groups were seen every
day in the 1250 m elevational zone. Average
group size in the reserve was 2.3 0.8 individ-
uals. Researchers observed individuals in the
1250 m zone eating Cryptocarya sp. the major
fruiting species in this zone other than Canarium.
We noted young individuals in 75% of these
groups.

Propithecus diadema candidus

Propithecus diadema candidus groups were ob-
served in the three highest elevational zones. Av-
erage group size in the reserve was 3.9 1.7
individuals. Young animals were observed in all
three zones, and 50% of the groups observed con-
tained young individuals. P. d. candidus groups
at the highest elevational zone came quite close
to the observers, enabling us to observe their pel-
age color well. Individuals in the same group had
coat colors that ranged from all white (the ex-
pected morph for the subspecies) to mostly white
with a black or yellow pygal patch to animals with
faint colors in a pattern that resembled P. d. dia-
dema pelage. Notable pelage characteristics: some
individuals were pure white; some had a dark
brown or yellow pygal patch; face black or pink;
thick white fur at underarm/triceps; two individ-
uals had black heads; one individual had a thick
black stripe from the top of the head down the
back and across the shoulders that lightened pos-
teriorly; one individual had light amber color on
upper portions of shoulder and thigh; and one in-
dividual had a thin black stripe down the back.

Species Accumulation Curves

Diurnal and nocturnal primate species were ob-
served along transects within 2 and 16 person-
hours of observation (Figure 13-1) across eleva-
tions. One species (Eulemur rubriventer) was ob-
served in the 450 m zone but was not observed
along the transects. Total effort for both nocturnal
and diurnal species was less than in previous stud-
ies because only one primatologist worked this
survey, both day and night. In addition, at the
highest elevational zone, the existing trails were
surveyed at least once a day, but there was very
little forest and the trails were quite short (Tables
13-3 and 13-4), so it was difficult to get quanti-
tative results in a short time.

Discussion

The total number of lemur species found in the
PN de Marojejy (10) is about average for eastern
rainforest primate communities. The closest rain-
forests for which recent survey information is
available are the RS d'Anjanaharibe-Sud, where
1 1 species were observed (Schmid & Smolker,
1998), and the PN de Masoala, where 10 species
were seen (Table 13-5). Species composition var-
ied among these sites, with some species observed
in only one of the three sites (e.g., Indri indri in
the RS d'Anjanaharibe-Sud and Varecia variegata
rubra in the PN de Masoala). The PN de Masoala
has a comparatively impoverished diurnal primate
fauna of only four taxa. It is still unclear why
Indri indri is found 20 km to the southwest in
Anjanaharibe-Sud (Schmid & Smolker, 1998) and
not in Marojejy.

At the lowest elevational zone of the PN de
Marojejy, we heard two species that we did not
expect to find. Outside Marojejy, Eulemur rubri-
venter has mainly been found at middle elevations
and above. This is the first published record of E.
rubriventer at low altitudes. They are probably
present at low densities at this altitude. Phaner
had not been observed previously in the PN de
Marojejy, but eastern populations of this species
are usually found in low density, and individuals
seem to vocalize less often than those in western
populations. If the individuals heard were indeed
Phaner, it would not be too surprising, as their
distribution includes areas to the north (Daraina)
and south (Masoala Peninsula) of PN de Marojejy
(Mittermeier et al., 1994). No Allocebus trichotis

STERLING AND MCFADDEN: LEMURS

271

450 m

775 m

t 1 r

8 10 12 14

Person hours

Person hours

1250 m

1625 m

t i i i i r
2 4 6 8 101214161820

Person hours

"o
a>
o.

E
z

on
4-
3-

2-

1 -
0-C

7 i i

i

i

8 10

Person hours

1875 m

Nocturnal species
Diurnal species

12 3

Person hours

Fig. 13-1. Species accumulation curves for primate populations. October-November 1996. at five sites in the PN
de Marojejy.

was observed during the course of this study, de-
spite their having been found in RS d'Anjana-
haribe-Sud. to the southwest (Schmid & Smolker,
1998).

The fact that a Microcebus rufus infant was
found with its eyes open in mid-October is un-
usual, given that M. rufus populations in the PN
de Ranomafana mate in mid-September, with
births in November (Atsalis, 1998). Gestation in
this species is thought to last 60 days, and infants
open their eyes within 2-4 days of birth (Glatston,

1979), which puts conception at mid- August. Re-
production in mouse lemurs could be affected by
a variety of factors, including temperature, which
itself might vary at any one time between alti-
tudes.

Experience from previous multitaxa expedi-
tions has suggested that the line transect method
may not always be the optimal method for esti-
mation of primate density in short surveys be-
cause low numbers of sightings do not allow for
the determination of the area surveyed (Sterling

272

FTELDIANA: ZOOLOGY

Table 13-5. Species observed in the PN de Marojejy and neighboring protected areas.

Species

PN de Marojejy

RS d'Anjanaharibe-Sud*

PN de Masoalat

Eulemur fulvus albifrons

X

X

X

Eulemur rubriventer

X

X

Hapalemur griseus griseus

X

X

X

Indri indri

X

Propithecus diadema candidus

X

X

Varecia variegata rubra

X

Avahi laniger

X

X

X

Lepilemur mustelinus

X

X

X

Allocebus trichotis

X

X

Microcebus rufus

X

X

X

Cheirogaleus major

X

X

X

Phaner furcifer

X

X

Daubentonia madagascariensis

X

X

X

Total species

10

11

10

* Schmid and Smolker, 1998.

f Sterling and Rakotoarison, 1998.

& Ramaroson, 1996; Duckworth, 1998; Sterling
& Rakotoarison, 1998). This was the case in this
particular study, as the total number of observa-
tions of lemur groups or individuals, diurnal or
nocturnal, was low. The fact that Eulemur rubri-
venter was recorded on audiotape, but not ob-
served, underscores the inadequacy of 10 days or
less of surveys for estimation of primate richness
and density by one experienced researchers in an
area. Thus, it is difficult to analyze or compare
the density data. Although density estimates are
given for both methods (for the line transect meth-
od, we only calculated relative density estimates),
we consider these data preliminary and subject to
further research. The experimental method seems
to be feasible from a logistical standpoint for col-
lecting data on diurnal primates during short sur-
veys, but it must be tested in an area of known
density, to assess its accuracy. This method would
be susceptible to differences in observer reliabil-
ity, the quality of home range size data from other
studies, and number of person-hours spent in the
survey zone. The low number of total observa-
tions per species at each elevation limited the ef-
ficacy of this method during the current survey.
These low numbers were probably related to cli-
mate, as the weather was unseasonably cold and
wet during the survey.

Hunting pressure on lemurs on the lower slopes
of the southeastern portion of the reserve appears
to be relatively high, given both the reaction of
diurnal groups to the presence of humans and the
presence of lemur traps. Adequate conservation of
this reserve is critical to the survival of Propithe-
cus diadema candidus, as it is only one of two

existing protected areas (along with RS
d'Anjanaharibe-Sud) that harbor this endangered
subspecies.

Acknowledgments

We thank the Center for Biodiversity and Con-
servation at the American Museum of Natural
History and the World Wide Fund for Nature for
their financial and technical support for this re-
search. We also thank Steven Goodman, Magi Ra-
belaza, Faratiana Ranaivojoana, Jean Claude Ra-
zafimahaimodison, and Frank Hawkins for their
assistance in data collection.

Literature Cited

Atsalis, S. 1998. Feeding ecology and aspects of life-
history in Microcebus rufus (family Cheirogaleidae,
order Primates). Ph.D. diss., City University of New
York, New York, NY.

Benson, C. W., J. F. R. Colebrook-Robjent, and A.
Williams. 1976. Contribution a l'ornithologie de
Madagascar. Oiseaux et R.F.O., 46: 103-134, 209-
242, 367-386.

1977. Contribution a l'ornithologie de Mada-

gascar. Oiseaux et R.F.O., 47: 41-64, 167-191.

Duckworth, J. W 1993. Feeding damage left in bam-
boos, probably by aye-ayes {Daubentonia madagas-
cariensis). Folia Primatologica, 14: 927-931.

. 1998. The difficulty of estimating population

densities of nocturnal forest mammals from transect
counts of animals. Journal of Zoology, 264: 466-468.

Glatson, A. R. H. 1979. Reproduction and behavior of

STERLING AND MCFADDEN: LEMURS

273

the lesser mouse lemur (Microcebus murinus. Miller
1777) in captivity. Ph.D. diss. University of London.
University College. London. United Kingdom.

Goodman. S. M.. and E. J. Sterling. 1996. The utili-
zation of Canarium (Burseraceae) seeds by vertebrates
in the RNI d'Andringitra. Madagascar, pp. 83-89. In
Goodman, S. M.. ed.. A floral and faunal inventory
of the eastern slopes of the Reserve Naturelle Inte-
grate d'Andringitra. Madagascar: With reference to
elevational variation. Fieldiana: Zoologv. n.s.. 85: 83-
89.

GU1LLAUMET, J. L.. J. M. Betsch. C. Blanc. R Morat.
A. Peyrieras. sous la direction de R. Pal li an.
1975. Etude des ecosystemes montagnards dans la re-
gion malgache. III. Le Marojezy. IV. Lltremo et
lTbity. Geomorphologie. climatologie. faune et flore
(Campagne RCP 225, 1972-73). Bulletin du Museum
National d'Histoire Naturelle (Paris), serie 3. 309
(Ecologie Generale). 25: 27-67.

Humbert. H. 1955. Une merveille de la nature a Mad-
agascar. Premiere exploration botanique du massif du
Marojejy. et de ses satellites. Memoires de l'lnstitut
Scientifique de Madagascar, serie B. 6: 1-210.

Iwano. T.. and C. Iwakawa. 1988. Feeding behavior of
the aye-aye (Daubentonia madagascariensis) on the
nuts of ramy {Canarium madagascariensis). Folia Pri-
matologica. 50: 136-142.

Merenlender. A.. C. Kremen. M. Rakotondratsima.
and A. Weiss. 1998. Monitoring impacts of natural
resource extraction on lemurs on the Masoala Penin-
sula. Madagascar. Conservation Ecology, 2: article 5.
Available: www.consecol.ora/Journal.vol2/iss2/art5/

MlTTERMEIER. R.. I. TaTTERSALL W. R. KONSTANT, D.

M. Meyers, and R. B. Mast. 1994. Lemurs of Mad-
agascar. Conservation International. Washington. D.C.

Nicoll. M. E.. and O. Langrand. 1989. Madagascar:
Revue de la conservation et des aires protegees. World
Wide Fund for Nature. Gland. Switzerland. 374 pp.

Safford. R.. and W Duckworth. 1990. A wildlife sur-
vey of Marojejy Nature Reserve. Madagascar. ICBP.
Cambridge.

Schmid. J., and R. Smolker. 1998. Lemurs of the Re-
serve Speciale d*Anjanaharibe-Sud. Madagascar, pp.
227-238. In Goodman. S. M.. ed.. A floral and faunal
inventory of the Reserve Speciale d'Anjanaharibe-
Sud. Madagascar: With reference to elevational vari-
ation. Fieldiana: Zoology, n.s.. 90: 227-238.

Sterling. E. J. 1993. The behavioral ecology of the
Aye-aye on Nosy Mangabe. Madagascar. Ph.D. Diss..
Yale University. New Haven. Conn.

Sterling. E. J., and M. G. R\maroson. 1996. Rapid
assessment of the primate fauna of the eastern slopes
of the Reserve Naturelle Integrale d'Andringitra.
Madagascar, pp. 293-305. In Goodman, S. M.. ed.. A
floral and faunal inventory of the eastern slopes of the
Reserve Naturelle Integrale d'Andringitra. Madagas-
car: With reference to elevational variation. Fieldiana:
Zoologv. n.s.. 85: 293-305.

Sterling. E. J., and N. Rakotoarison. 1998. Rapid as-
sessment of the primate species richness and density
on the Masoala Peninsula, eastern Madagascar. Folia
Primatologica. 69: (suppl. 1): 109-116.

Tattersall. I. 1982. The primates of Madagascar. Co-
lumbia University Press. New York.

274

FIELDIANA: ZOOLOGY

Gazetteer of Localities Mentioned in the Text*

Longitude

South

Latitude East

Elevation

Locality

()

(')

(')

(m)

Alaotra, Lac

48

30

17

30

Ambalamanasy

49

36

14

32

Ambanizana

49

58

15

37

Ambatoharanana

48

45

14

26

Ambatosoratra, Mt.

49

42

14

33

1583

Ambatovaky, RS

-49

15

-16

42

Ambavaomby River

49

46

14

26

Ambinanitelo River

49

47

14

26

Ambodisovoka

48

17

20

05

Ambohimitambo (see Ambohimitombo)

Ambohimitombo

47

23

20

43

Ambohitantely, RS

-47

16

-18

09

1450-1660

Ampahana

50

13

14

45

Ampanasankolana

49

46.5

14

26.2

300-525

Ampanihy

44

45

24

42

Ampitambe*

47

46

20

22

Ampombilava

48

15

13

24

Analamazaotra, RS

48

25

18

56

930-1040

Andampimbazaha River

49

45.7

14

26.0

775

Andapa

49

39

14

40

Andasibe (see Analamazaotra)

Andohahela, PN (parcel 1)

-46

11

-24

42

Andranofototra River

49

46

14

28

Andranomay Forest

47

57

18

29

1300

Andringitra, PN

-46

54

-22

14

700-2658

Androranga River

49

46

14

18

Anjanaharibe-Anivo

49

27

14

44

2064

Anjanaharibe-Sud, RS

-49

26

-14

42

500-2064

Anjenabe, Mt.

49

44

14

18

Ankafana (see Ankafina)

Ankafina

47

12

21

12

Ankarafantsika, RNI

-46

57

-16

09

80-330

Ankaratra Massif

47

12

19

25

2642

Anjozorobe

47

52

18

24

Antalaha

50

16

14

53

Antanana

47

31

18

55

Antongil, Baie d'

49

50

15

45

Antranohofa

49

44.5

14

26.0

Antsahabe River

49

54

14

30

Antsahaberoakahely River

49

39

14

23

Antsiranana

49

17

12

16

Bedinta (see Ampanasankolana)

Behondroko, Mt.

49

46

14

24

Belaoko

49

44

14

34

Beondroka, Mt. (see Behondroko)

Berenty, RP

46

17

24

59

Betaolana

49

27

14

31

Betsomanga River

49

45

14

17

Camp 1 (1996)

49

46.5

14

26.2

450

Camp 2 (1995)

49

45.7

14

26.0

775

Camp 3 (1995)

49

44.5

14

26.2

1225

Camp 4 (1995)

49

44.5

14

26.4

1625

Camp 5 (1995)

49

44.1

14

26.8

1875

Daraina

49

40

13

12

Didy

48

32

18

07

Diego-Suarez (see Antsiranana)

Djabal River (see Djabala River)

Djabala River

48

13

13

23

Doany

49

31

14

22

Gazetteer

275

Longitude South

Latitude East

Locality

()

(')

o

(')

Elevation

(m)

Fanovana

Farafangana

Farahalana

Fito

Fort-Dauphin (see Tolagnaro)

lharana (see Vohemar)

Ikongo Forest

Ivohibe, RS

Kirindi Forest

Lakato

Lohan Andranomifotrotra

Lokoho River

Mahajanga

Mahanara River

Maimborondro (see Mainborondro)

Mainborondro, Mt.

Maintirano

Majunga (see Mahajanga)

Mananano River

Mananara, RB

Mananara River

Mananjary

Manantenina (village)

Manantenina River

Mandena

Manjakatompo

Manombo, RS

Manongarivo, RS

Mantadia (see Mantady)

Mantady, PN

Maroambihy

Maroantsetra

Marojejy, PN

Marojezy (see Marojejy)

Marolambo

Marosse, Isle (see Nosy Mangabe)

Marovoay

Masoala, PN

Midongy du Sud

Montagne d'Ambre, PN

Morondava

Namorona River

Nandihizana

Nosivolo River

Nossi-Be (see Nosy Be)

Nosy Be

Nosy Mangabe

Perinet (see Analamazaotra)

Ranomafana, PN

Ranomafana (Ifanadiana)

Ranomena

Rantabe

Sahasinaka

Sambava

Sihanaka Forest

Tamatave (see Toamasina)

Tampolo

Toamasina

Tolagnaro

Tolongoina

Tsaratanana, RNI

48

34

47

50

50

10

48

54

43

33

-46

59

44

43

48

22

49

44.1

49

44

46

19

49

58

49

47

44

01

48

03

-49

42

46

33

48

20

49

49

49

49

49

49

47

26

47

44

-48

23

-49

27

49

30

49

44

-49

15

48

07

46

39

-50

10

-47

01

-49

10

44

17

48

13

-47

10

48

30

48

15

49

46

-47

28

47

28

47

17

49

39

47

49

50

10

48

32

49

25

49

23

46

59

47

31

48

51

18

55

22

49

14

26

18

05

21

47

-22

32

20

03

19

03

14

26.4

14

37

15

43

13

58

14

25

18

03

22

02

-16

22

24

50

21

14

14

30

14

29

14

29

19

22

23

02

-13

59

-18

51

14

30

15

26

-14

26

20

02

16

06

-15

38

-23

35

-12

37

20

17

21

39

-20

50

19

56

13

20

15

30

21

16

21

14

23

25

15

42

21

49

14

16

18

07

17

17

18

10

25

01

21

33

13

59

775-2060

1050
2133

119

0-570

0-137
150-1876

75-2133

850-1357

0-40
227-2876

276

FIELDIANA: ZOOLOGY

Locality

Vinanitelo
Vohemar
Vondrozo
Zahamena, RNI
Zombitse, PN

Longitude South

Latitude East

Elevation

() (')

()

(')

(m)

47 16

21

43

50 01

13

21

47 20

22

49

-48 50

-17

40

750-1512

-44 40

-22

47

485-825

* For geographical localities such as rivers, large reserves, and mountain ranges, we have given an intersection of
coordinates that allows for easy location on maps. Information presented in the Gazetteer is based partially on
coordinates given by the U.S. Board on Geographic Names (1955) and Viette (1991). PN = Pare National, RNI =
Reserve Naturelle Integrate, RB = Reserve Biosphere, RS = Reserve Speciale.

t Based on Carleton and Schmidt (1990).

Literature Cited

U.S. Board on Geographic Names. 1955. Madagascar,
Carleton, M. D., and D. F. Schmidt. 1990. Systematic Reunion and the Comoro Islands. Gazetteer 2. U.S.

studies of Madagascar's endemic rodents (Muroidea: Government Printing Office, Washington, D.C.

Nesomyinae): An annotated gazetteer of collecting lo- Viette, P. 1991. Principales localites ou des Insectes ont
calities of known forms. American Museum Novita-"" et recueillis a Madagascar. Faune de Madagascar.

tes, 2987: 1-36. supplement 2. Private printing.

Gazetteer 277

Index to Scientific Names

Acanthaceae 46, 49, 51, 61, 65,
68, 69, 71, 75, 77, 79, 80, 95,
97
Acari 137, 138-140
Acariformes 138-140
Accipiter

francesii 180, 183, 184, 185

henstii 180, 186, 192, 196, 197

madagascariensis 183, 196, 197
Acridotheres

tristis 196
Acrocephalus

newtoni 181
Aframomum 10, 190, 191

angustifolium 35, 46, 65, 94
Afrocypholaelaps

ranomafanaensis 141
Agapornis

carta 180
Agauria 55, 99

salicifolia 47, 77, 79, 84
Agavaceae 80, 95
Agelaea

pentagyna 61, 63, 83
Agonostomus 148-149

catalai 148, 149

monticola 148

telfairii 148-149
Allberta 55, 79, 91

minor 75, 91, 103

sambiranensis 103
Albizia 45, 46, 60, 61, 63, 64, 65,

85
Alee do

vintsioides 180, 183, 184, 185
Alchornea 99
Alectroenas 189, 190

madagascariensis 180, 186, 189
A listrophoroides 1 3 8- 1 40
Alloc ebus

trichotis 266, 271, 273
Allophylus 69, 71, 93
Alluaudina

bellyi 162, 168, 171

Aloe 129

Amauropelta 24, 34

bergiana 24, 34
Ameroseiidae 137, 140
Amiota 129
Amphiglossus

astrolabi 161, 168

intermedius 161, 163, 168

macrocercus 161, 168, 171

mandokava 161, 168

melanopleura 161, 163, 168,
171

melanurus 161, 163, 168, 171

minutus 161, 163, 168, 171

mouroundavae 161, 163, 168,
171

ornaticeps 161, 168, 171

punctatus 161, 163, 168, 171
Anacardiaceae 67, 80, 95, 97

Analgidae 140
Androlaelaps 141
Angiopteris 65, 90

madagascariensis 24, 34
Angraecum 71, 73, 89, 102
Angw///a

bengalensis
labiata 147

bicolor 147

marmorata 147

mossambica 147-148
Anguillidae 147-148
Anhinga

rufa 180
Anisophyllea

fal lax 61, 80
Anisophylleaceae 80, 95
Annonaceae 49, 51, 60, 61, 62, 65,

67, 80, 95, 97
Anodonthyla

boulengeri 166
Anoplura 139-140
Anf/ioc/e/sta

longifolia 67, 87

madagascariensis 73, 75, 87,
100
Antidesma 61, 85, 99, 100

petiolare 69, 85
An?ro/?/rvwm

boryanum 24

malgassicum 24
Apenthecia 129
Ap/j/oia

theiformis 69, 73, 75, 77, 79,
85, 100
Apiaceae 79, 80, 95, 97
Apocynaceae 49, 51, 65, 80, 95,

97
Apodocephala 71, 81, 97
Appolonias 100

A/7MS

barbatus 180, 185

me/fca 180
Aquifoliaceae 81, 95, 97
Araceae 81, 95
Araliaceae 49, 51, 55, 75, 79, 81,

95, 97
Araneae
Ardea

purpurea 180
Ardeola

idae 180

ralloides 180
Arecaceae49, 51, 81, 95, 97
Aristolochia 69, 81
Aristolochiaceae 81, 95
Artabotrys 61, 63, 65, 67, 80
Arthropteris

monocarpa 24, 34

orientalis

var. subbiaurita 24

palisotii 24, 29
Arundinaria 75, 90

Ascarina 98
Asclepiadaceae 97
Asi'o

madagascariensis 180, 185, 270
Asplenium 19, 20, 24, 28, 32, 33,
38, 73, 90

o#m<- 24, 33

a/ze//// 24

andapense 20, 29

bipartitum 24, 34

cancellatum 24, 34

cuneatum 24, 34

dregeanum 24, 34

friesiorum 24

herpetopteris

var. acutipinnatum 24, 33
var. herpetopteris 24
var. villosum 24

inaequilaterale 24

marojejyense 20, 24, 29, 34

mVtos 24, 34, 45, 61, 63, 65, 69,
90

normale 24

obscurum 24, 33

pellucidum 24

petiolulatum 24

sandersonii 24, 36

thunbergii 24

unilaterale 24, 34

var. paucijugum 24, 34
Asteraceae 47, 49, 51, 55, 67, 69,
71, 73, 75, 77, 79, 81-82, 95,
97-98
Ate/orai's

crossleyi 180, 186, 193, 196,

197
pittoides 180, 186, 192, 193,
196, 197
Atemnosiphon 75, 94

coriaceus 104
Athyrium
scandicinum

var. bipinnata 24, 33
var. scandicinum 24
Atopomelidae 138-140
Ara/?/

/an/ger 265, 267, 268-269, 270,
273
Avenzoariidae 140
Av/raia

madagascariensis 180, 185, 192

AvV0fOM5

aeneofuscus 151

Bakerella 11, 87, 100-101, 190
Balsaminaceae 49, 51, 82, 95, 98
Beckeriella 105-112

fasciatus 107-109

Ji/ipirui 105, 107

maculata 108-111, 112
B^or/a 143, 144

longianalis \AA

INDEX

279

madagascariensis 144

marojejy 143, 144-147
Bedotiidae 143
Begonia 69, 73, 82, 98
Begoniaceae 82, 95, 98
Beguea 65, 69, 93
Beilschmiedia 67, 86
Belvisia

spicata 24
Bembicia 71, 85
Bernieria 182, 190, 191

cinereiceps 181, 184, 185, 190,
195, 196, 197

madagascariensis 181, 184, 185,
186, 190, 195

tenebrosa 175, 181, 183, 185,
188, 190, 194-195, 197

xanthophrys 181, 184, 185, 186,

190, 195, 196, 197
zosterops 140, 181, 184, 185,

186, 190, 191
Bertiera 61, 63, 91
Bignoniaceae 82, 95, 98
Blaesodactylus

antongilensis 161, 167
Blechnum 28, 32, 38, 79, 90

attenuatum 24, 34, 36

bakeri 24, 33

humbertii 20, 24, 29

ivohibense 24, 33

longepetiolatum 20, 24, 29, 34

madagascariense 24

simillimum 24
binerve 24

simillimum

var. xiphophyllum 24, 33
Blotiella

coriacea 20, 24, 29

madagascariensis 24, 36
Boa

manditra 162, 168, 171
Boehmeria 69, 94
Boidae 162, 168, 171
Bolbitis

auriculata 24, 34

humblotii 24, 33
Boophis 167, 172

albilabris 160, 161, 167, 170

anjanaharibeensis 160, 167, 170

brachychir 160, 167, 170

burgeri 164, 167

englaenderi 160, 167

goudoti 164

/wtews 160, 167, 170

madagascariensis 160, 167, 170

majori 167

marojezensis 160, 167, 170

rappioides 161, 167, 170

reticulars 161, 164, 167, 170

rufioculis 164

tephraeomystax 161, 167
Brachylaena 46, 69, 81

merana 75, 81, 97
Brachypteracias 191

leptosomus 180, 184, 186, 190,

191, 192-193, 196, 197

squamiger 180, 193, 196, 197
Brachytarsomys 252

albicauda 233, 251, 252, 258,
259
Brachyuromys 260

betsileoensis 259, 260

ramirohitra 259, 260
BnVte/za 61, 65, 67, 85, 100
Brochoneura

acuminata 45, 46, 60, 61, 63, 88

fcettc/i/ 161, 165, 167, 171
griveaudi 161, 163, 165, 167
forcfc*'/ 158, 161, 165, 167, 172
minima 161, 165, 167
superciliaris 164, 167
vaJon/ 161, 165, 167, 171

Bubulcus
ibis 183

Bulbophyllum 67, 73, 75, 79, 89-
90, 102

Burseraceae 49, 51, 82, 95, 98,
267

Buteo

brachypterus 180

Butorides
striatus 183

Byttneria 67, 93

Cacojce/iMS 129
Caesaria 85
Calanthes 102
Calicalicus 191

madagascariensis 181, 187, 191
Calodrosophila 130, 132
Calophyllum 47, 63, 67, 71, 73,

74, 75, 76, 77, 82-83, 98
drouhardi 98

Calumma
bifidus 167

boettgeri 161, 165, 167, 171
brevicornis 161, 165, 167, 169,

172
cucullata 161, 165, 167
gastrotaenia 158, 161, 165, 168,
169, 171
guillaumeti 165, 172
marojezensis 165
globifer 168
ma/r/ze 161, 168, 171
nasMta 161, 165, 168, 171
peyrierasi 161, 165, 168, 172
Campnosperma 75, 81, 97
Canarium 53, 98, 267, 270, 271
froiV/nii 46, 61, 63, 65, 66, 67,

82
madagascariense 61, 69, 82, 98
Canellaceae 82, 95, 98
Canephora 65, 91
Canirallus

kioloides 180, 186
Canthium 46, 67. 69, 70, 71, 73,

75, 77, 79, 91. 103
Caprimulgus

enarratus 183
madagascariensis 180, 183, 185

Carissa 63, 80

Casearia 69, 73, 75, 77, 100

Cassinopsis 47, 63, 65, 71, 75, 76,

77, 79, 86, 100
Cassipourea 71, 75, 91, 102
Celastraceae 73, 75, 79, 82, 95, 98
Centropus

toulou 180, 186
Chamaeleonidae 161, 165, 167,

171
Chapelieria 103
Cheilanthes 24, 29, 33
Cheirogaleus

major 265, 267, 269, 270, 273
Chernetidae 139
Chloranthaceae 77, 82, 95, 98
Chris tella

dentata 24

distans 24
Chrysophyllum 53

boivinianum 45, 46, 53, 60, 61,
62, 63, 64, 65, 93
Chymomyza 130, 132
Cichlidae 149-150
Cinnamosma 67, 82

madagascariensis 98
Cisticola

cherina 181
Citrus 104
Clerodendron 61, 71, 73, 75, 77,

79, 94, 104
Clusiaceae 49, 51, 53, 82-83, 95,

98-99
Cochlidium 21

serrulatum 24, 36
Colea 69, 82, 98
Colubridae 162, 168, 171
Coniogramme

madagascariensis 24
Connaraceae 83
Convolvulaceae 65, 83, 95
Copsychus

albospecularis 140, 181, 184,
186
Coracina

cinerea 180, 186
Coracopsis 180, 189, 190

nigra 181, 186

vasa 181
Cordylidae 161, 168, 171
Corvus

albus 183
Coturnix

coturnix 180, 183
Cotylopus

acutipennis 151
Coua 191

caerulea 180, 186, 191

cristata 180

reynaudii 180, 186

serriana 180, 186
Crossleyia 182, 190
Croton 65, 71, 85, 100
Cryptocarya 46, 55, 63, 64, 65,
66, 67, 69, 70, 71, 73, 86,
100, 271

280

FIELDIANA: ZOOLOGY

Cryptoprocta 270
Cryptosylvicola

randrianasoloi 181, 184, 185,
187, 194
Ctenitis 25, 33

ochrorachis
var. violacea 24

poolii 25

warburii 25, 33
Ctenopteris 29, 32, 33, 38

alboglandulosa 25

devoluta 25, 36

elastica 25

flabelliformis 21, 25

humbertii 25

villosissima 25

zenkeri 25, 36
Cuculus 191

rocA/i 180, 186, 191
Cunoniaceae 49, 51, 83, 95, 99
Cyanolanius

madagascarinus 181, 187
Cva?^a 28, 32, 33, 38, 46, 61, 65,
66, 68, 69, 71, 73, 77, 79, 90

albida 25, 33

alticola 25, 29

approximata 25

bellisqamata

var. basilobata 25, 34

boivinii 25

borbonica 25, 33

6w//ara 25, 36, 73, 75, 90
var. madagascarica 25, 33

costularis 25, 34, 61, 90

coursii 25

decrescens 25, 34, 46, 62, 63,
65, 71, 90

dregei 25

var. polyphlebia 25

hildebrandtii 65, 90

ligulata 25, 34

longipinnata 25, 33

melleri

var. melleri 25

perrieriana 25

serratifolia 25, 33
Cyatheaceae 95
Cyclosorus

interruptus 25, 33
Cynorchis 11, 90, 102
Cyperaceae 83, 95, 99
Cyperus 99
Cvpj/r5

/jarvws 180, 185

Danais61, 69, 71, 77, 91

tsaratananensis 103
Daubentonia

madagascariensis 265, 266, 267,
269-270, 273
Davallia

chaerophylloides 25, 34

repens 24, 34
Decarydendron 101
Deinbollia 45, 61, 65, 93
Demodex 139-140

Demodicidae 139-140
Deparia 25, 34

marojejyensis 20, 29

parvisora 25
Dettopsomyia 129
Deuteromallotus 46, 61, 63, 64,

65, 85
Dichaetanthera 71, 73, 75, 76, 77,

78, 79, 87, 101
Dichaetophora 69, 75, 130, 132
Dicranopteris

linearis 25
D/crMrw.s

forficatus 181, 187
Didymochlaena

microphylla 25, 34
Dillenia 67, 83

triquetra 67, 83
Dilleniaceae 83, 95
Dilobeia 53

thouarsii 53
Dionycha 101
Diospyros 45, 60, 61, 63, 65, 67,

69, 71, 73, 75, 83-84, 99
Diplazium 90

andapense 25, 34

dilatatum 25, 33

marojejyense 29

nemorale 25, 34

proliferum 25
Diptera 123, 125
Dolichopodidae 125
Dombeya 63, 65, 69, 71, 73, 93,

104
Dorstenia 101
Dracaena

re/teca 61, 65, 67, 69, 71, 77,
80
Dromaeocercus

brunneus 181, 186, 194, 196,
197
Dromicodryas

quadrilineatus 162, 168
Drosera 99
Droseraceae 99

Drosophila 124, 127, 128, 130,
132

busckii 133

funebris 133

/ryoV/ 133

immigrans 133

malerkotliana 133

melanogaster 124, 130, 133, 134

montium 130

simulans 133
Drosophilidae 123-134
Drynan'a

willdenowii 25, 33
Dryolimnas

cuvieri 183
Dryoptera

mangindranensis 25, 34
Dry petes 100

Dy/w/s 46, 47, 61, 63, 65, 66, 67,
68, 69, 70, 71, 72, 73, 74, 75,
77, 79, 81, 97, 270

Dyscophus
insularis 166

Ebenaceae 49, 51, 83-84, 95, 99
EbenaWa

inunguis 161, 167, 170
EgreWa
a/foa 180
dimorpha 180
Elaeocarpaceae 49, 51, 75, 84, 95,

99
Elaeocarpus 11,11, 84, 99
Elaeodendron 46, 69, 71, 75, 85
Elaphoglossum 26, 28, 29, 32, 33,
34, 36, 38, 69, 77, 79, 90
achroalepis 25
acrostichoides 25
angulatum 25
aubertii 25
cerussatum 20, 29
decaryanum 25
aecfcen//

var. rufidulum 25
forsythii-majoris 25, 33
humbertii 26
lepervanchii 26
leucolepis 26

marojejyense 20, 26, 29, 34
petiolatum

ssp. salicifolium 21, 26, 33
scolopendriforme 26
sieberi 26
spathulatum 26, 33
subsessile 26
Eleotridae 150-151
Eleotris 150
/wsca 150-151
melanosoma 150, 151
Eliurus 233, 251
grandidieri 137, 139, 141, 231,
233, 242, 243, 244, 245-
247, 250, 251, 254, 255,
256, 257, 258, 259, 260
mayon 139, 219, 231, 235-242,
243, 244, 246, 247-248,
252, 254, 255, 256, 257,
258, 259, 260, 262-263
m/nor 139, 219, 231, 233, 242,
243, 244, 246, 248, 249,
254, 255, 256, 258, 259
myoxinus 233, 248, 251
penicillatus 231, 232, 235-238,

239, 242, 263
tana/a 231, 241, 242, 243, 244,
246, 248, 249, 255, 256,

258, 259

r>?w 139, 231, 232, 238-242,
243, 244, 246, 248-250,
254, 255, 256, 257, 258,

259, 260, 263
mfc<?//a 77, 79, 88, 101
Empididae 125
Enteromorpha 97
Enterospermum 67, 69, 71, 91-92
Ephippiandra 46, 53, 63, 65, 68,

69, 88

INDEX

281

madagascariensis 101
perrieri 101
Ephydridae 105
Ephydra

bispinosa 106
Ereynetidae 140
Erica 47, 54, 55, 75, 77, 79, 84,

99
Ericaceae 47, 49. 51, 55, 84. 95,

99, 260
Erima 129
Eriocauleaceae 99
Eriocaulus 99
Erythroxylaceae 84. 95. 99
Erxthroxxlum 63. 67, 69. 73. 75.

79. 84, 99
Eugenia 46, 47. 61. 63, 65. 67. 69.
70. 71, 72, 73. 75, 77. 79, 89.
102
emirnensis 75. 89
Eulemur
fulvus 268

albifrons 265, 267, 268, 270-
271, 273
rubriventer 265, 267, 268. 270.
271, 273
Eulistrophoroides
Evodia 13, 104
Euphorbiaceae 49, 51. 67. 75, 85,

95, 99-100
Eurxceros 191
prevostii 181, 184, 185, 187,
191, 195. 196, 197
Eurylaimidae 140
Eurxstomus

glaucurus 180. 183, 186
Eutriorchis

astur 178, 183, 192, 196, 197
Exacum 100

Fabaceae49. 51. 85. 95, 100
Falco

concolor 183

eleonorae 180. 183

newtoni 180

peregrinus 180, 183

zoniventris 183
Faucherea 75. 93, 104

hexandra 67, 93

parvifolia 71, 93

thouvenotii 104
Faurea

forficuliflora

var. elliptica 102
Ficus 65. 69. 88, 101, 189

lutea 63, 65. 88

politoria 101

soroceoides 69, 88

tiliaefolia 101
Filicium 71. 73, 93
Flacourtiaceae 49. 51. 63. 73, 75,

85-86, 95. 100
Fluggea 100
Foudia

madagascariensis 181. 184, 187,
191

omissa 140. 181. 184. 187
Furcifer
pardalis 161. 165, 168, 171

Gaertnera 61, 69, 71, 73. 75. 77.
79, 92. 103

macrostipula 71, 75. 92, 103

uniflora 92, 103
Gastropini 106
Gastrops 106
Geckolepis

maculata 161, 167, 170
Gekkonidae 161. 167. 170
Gentianaceae 100
Geodipsas 162. 168, 172

heimi

infralineata 168

laphystia 162. 168
Gerrhosauridae
Gesneriaceae 100
Gitona 129
Glecheniaceae 79, 91
Gleichenia

madagascariensis 26, 34

polxpodioides 26
Glycyphagidae 138-140
Gobiidae 151-153
Grammitidaceae 21
Grammitis 19. 20, 26, 28, 32, 33,
34, 36, 38

barbatula 26. 33

copelandii 26, 34

cryptophlebia 26, 34

ebenina 26

gilpinae 26

holophlebia 26

kyimbilensis 26

microglossa 26. 33

obtusa 26

sxnsora 26
Gravesia 61. 67. 69, 71. 73. 87
G^H'/a 65, 67, 71. 73. 94, 104
Gymnomyzinae 106
Gxm?ruromxs

roberti 231, 233, 242, 243, 244,
246, 250, 255. 256. 258.
259

Haemaphysalis 1 39
Hapalemur

griseus 268, 270
griseus 265, 267, 268, 270,
273
Hartertula

flavoviridis 183. 197
Harllaubius

auratus 181. 183
Helichrxswn 55. 75, 77. 79. 81-
82. 97-98

adhaerens 79. 81, 97
Hemicentetes 204, 21 1. 221. 228

nigriceps 211, 221

semispinosus 220. 221
Heterixalus

betsileo 162, 166

madagascariensis 160, 166

Hirtodrosophila 127, 129
Histiopteris

incisa 26
Homalium 63. 67, 73, 77, 85
Homopholis

antongilensis
Hugonia 67, 87
Humata

repens 25
Huper-Ja 29, 32. 38

cavifolia 26

megastachxa 26

obtusifolia 26. 33

ophioglossoides 26

pecten 26

saururus 33

squarrosa 26

suberecta 26

trigona 26

verticillata 26
Hxmenophxllum 28, 32, 33, 36, 38

capense 26

capillare 26

deltoideum 26

fumarioides 26, 34

heimii 26

hirsutum 26

humbertii 26

inaequale 26

panum 26

polyanthos 26

poolii 26

sibthorpioides 26, 34

tunbrigense 26

veronicoides 27

viguieri 27
Hymenoptera 125
Hyperacanthus 103
Hyperolidae 160. 166
Hypositta

corallirostris 183
Hypsipetes

madagascariensis 180, 184, 185.
186

Icacinaceae 75. 86, 95, 100

mm* 61. 69. 73, 81. 97
Impatiens 73, 75, 77, 79, 82, 98

mJn 271. 273
Insectivora
fco/o/ia 67. 80, 97
Ispidina

madagascariensis 180. 184. 185
Ithycyphus

blanci 168

miniatus 162. 168

Ixodes 139
Ixodidae 138-140
/.vora 71. 73. 92. 103

Jennealopsis 129
Juncaceae 100

282

FIELDIANA: ZOOLOGY

Labramia 69, 93
Laelapidae 138-140
Landolphia 61, 63, 65, 71, 80, 97
Lasteropsis 21

subsimilis 27, 33
Lauraceae 46, 49, 51, 53, 55, 62,
63, 70, 71, 75, 77, 86-87, 95,
100
Laurentomantis

horrida 162, 166

malagasia 160, 166
Leioheterodon

madagascariensis 162, 168
Lemyrea 103
Lepilemur 267

mustilenus 267, 269, 270, 273
Lepisorus

excavatus 27
Leptogale

gracilis 213
Leptopterus 191

c/w&erf 181, 187, 191

viridis 181, 187, 191
Leptosomus

discolor 180, 186
Leucophenga 129, 130
Liliaceae 63, 75, 87, 95
Linaceae 95

Lindsaea 19, 20, 27, 28, 29, 32,
36, 38

blotiana 27

coursii 27

ensifolia 27, 33

flabellifolia 27

goudotiana 27

madagascariensis 27

millefolium 27, 34

odontolabia 27, 33

odorata 27, 33
Lindsaeaceae 21
Liodrosophila 130, 132
Liophidium

rhodogaster 162, 168, 171

torquatus 162, 168
Liopholidophis 162, 168, 172

epistibes 162, 168, 171

.m<m#z 162, 168
Lipotyphla 201-229
Lissocephala 129
Listrophoroides 138-140
Loganiaceae 87, 95, 100
Lomariopsis

crass ifolia 27, 34
Lonchura 191

nana 181, 182, 187, 191
Lophotibis

cristata 180, 185, 191-192, 197
Loranthaceae 87, 95, 100-101,

190
Loxogramme

humblotii 27

lanceolata 27
Li/a7a

madagascariensis 71, 85
Luzonimyia 129

arctifasciatus 162, 168, 171

betsileanus 162, 168, 171

gaimardi 162, 168
Lycopodiella

caroliniana 21, 27, 33

cernua 27
Lycopodium 79, 91

zanclophyllum 27
Lygodactylus

miops 161, 167, 170
Lygodium

lanceolatum 27

Mafrifya

gravenhorstii 161, 168, 171
Macaranga 46, 62, 63, 65, 67, 69,
71, 73, 85, 100

cuspidata 65, 85, 100
Macrothelypteris

torresiana 27, 33
Madamyobia 138
Madlistrophoroides 138-140
Mflfl 69, 88, 101
Malleastrum 61, 63, 73, 88, 101
Mallophaga 140
Mallotus 46, 62, 63, 85, 100
Mammea 46, 61, 63, 65, 66, 67,
68, 69, 71, 73, 75, 77, 83, 98

bongo 65, 83
Mantella 160, 166, 172

laevigata 160, 166

madagascariensis 166

nigricans 160, 166
Mantellidae 160, 166, 170
Mantidactylus 160, 167, 172

ag/avei 160, 167, 170

albofrenatus 160, 166, 170

albolineatus 166

as/w 160, 166, 170

betsileanus 160, 166, 170

bicalcaratus 160, 166, 170, 172

fo//?o/T 160, 166, 170

cornutus 160, 166, 170

cHrto 160, 166, 170

femoralis 160, 166, 170

flavobrunneus 166

grandidieri 160, 166, 170

grandisonae 166, 170

granulatus 166

guttulatus 166

klemmeri 160, 166, 170

leucomaculatus 160, 166

fifcer 160, 166, 170

lugubris 160, 166, 170

/wtews 160, 166, 170

majori 162, 167

opiparis 160, 166, 170

peraccae 160, 167, 170

pliciferus 167

pseudoasper 162, 167

/jw/c/ier 160, 167, 170, 172

punctatus 167

redimitus 160, 167, 170

rrWco/a 160, 167, 170

spiniferus 162, 167

ulcerosus 160, 167, 170

Mapouria 61, 63, 71, 73, 79, 92

pyrrotricha 103
Maranthaceae 61, 87, 95
Mara#/a

fraxinea 27
Margaroperdix

madagascariensis 180, 183
Marojejya 11
Mauloutchia

humblotii 53
Medinilla 61, 65, 69, 71, 73, 75,

77, 79, 87
Melanophylla

humbertiana 69, 71, 87, 101
Melanophyllaceae 87, 95, 101
Melastomataceae 49, 51, 63, 67,

73, 75, 77, 87-88, 95, 101
Meleastrum 271
Meliaceae 88, 95, 100
Memecylon 65, 69, 71, 73, 87, 101
Mendoncia 71, 80
Menispermaceae 95
Merops

superciliosus 180
Mesitornis

unicolor 180, 183, 185, 192,
196, 197
Microcebus

rufus 265, 267, 269, 270, 272,
273
Microdrosophila 131, 132
Microgale 201, 202, 203, 209,
213, 219, 222, 223, 224, 225,
226, 228

brevicaudata 201, 202, 203,

206, 207, 208-210, 211,
212, 222, 225, 229

breviceps 206, 210
cowani 138, 201, 202, 203, 207,
208, 209, 210, 211,212,

219, 222, 223, 225, 229,
243

dobsoni 138, 201, 202, 203,

208, 209, 210-213, 218,

220, 221, 222, 223, 225,
229

drouhardi 211

dryas 201, 202, 211, 221, 222,

223
fotsifotsy 201, 202, 203, 207,

209, 211, 212, 213, 220,

221, 222, 225, 228
gracilis 201, 202, 203, 207, 209,

211, 213-214, 219, 221,

222, 223, 229, 243
gymnorhyncha 138, 201, 202,

203, 207, 209, 211, 212,
214, 221, 222, 223, 225,
229

longicaudata 138, 201, 202,
203, 207, 209, 211,212,
214-215, 222, 223, 225,
228

majori 214

monticola 138, 201, 202, 203,

207, 209, 211, 212, 215-

INDEX

283

216, 222, 223, 225, 226,

229
parvula 138, 201, 202, 203, 207,

209, 211, 212, 216, 221,

222, 223, 225, 228
principula 139, 201, 202, 203,

207, 209, 211, 212, 215,

216-217, 222, 225, 228
pulla 216
pusilla 220, 221
sorella 216
soricoides 139, 201, 202, 203,

207, 209, 211, 212, 217-

218, 222, 223, 225, 229
taiva 211, 221

talazaci 139, 201, 202, 203,

208, 209, 211, 212, 218,

219, 221, 222, 223, 225,
229, 243

thomasi 211, 216
Microgalobia 138-139
Microhylidae 160, 166, 170
Microlepia

madagascariensis 27
Microscalabotes

bivittis 161, 167, 172
Microsorum

punctatum 27
Milvus

migrans 183
Mirafra

nova 180, 183, 191
Monanthotaxis 97
Monimiaceae 49, 51, 53, 61, 77,

88, 95, 101
Monogramma

graminea 27
Monoporus 75, 79, 88, 101
Monti co la 182

sharpei 140, 181, 184, 185, 186,
196, 197
Monticolomys

koopmani 259
Moraceae 49, 51, 61, 63, 88, 95,

101
Morinda 103
Motacilla

flaviventris 180
Mugilidae 148
Muridae 139, 231-263
Murinae 231, 232, 242-245, 255,

258
Mus

musculus 259
Musaceae
Mussaenda

erectiloba 103
Mxcodrosophila 1 3 1
Myobiidae 138-139
Myrica 75, 79, 101

phillyreaefolia 1 1

spathulata 101
Myricaceae 88, 95, 101
Myristicaceae 49, 51, 88, 95
Myrsinaceae 47, 49, 51, 75, 79,
88-89, 95, 101-102

Myrtaceae 49, 51, 55, 89, 95, 102
Mystacornis

crossleyi 181, 187

Nastus 46, 47, 61, 63, 67, 69, 70,
71, 72, 73, 74, 75, 77, 78, 79,
90
Nectarinia

notata 181, 187
souimanga 181, 184, 185, 187,
188
Nematostylis 79, 92

anthophylla 103
Neodrepanis 193

coruscans 140, 180, 184, 185,

186, 190, 193
hypoxantha 137, 140, 180, 184,

185, 186, 190, 193, 196,
197

Neomixis 190

striatigula 181, 187, 190

tenella 181, 186, 190

viridis 181, 186, 190
Neotanygastrella 131, 132
Nephrolepis 63, 91

biserrata 27, 34

tuberosa 27, 33
Nesillas

typica 181, 184, 185, 186
Nesogale

dobsoni 210

talazaci 218
Nesomyinae 231, 243, 245-251,

255, 258, 259
Nesomys 231, 232, 250, 252

audebertii 252

rufus 231, 232, 233, 249, 250,
251, 252, 255, 256, 258,
259, 260
Nestis

cyprinoides 148

dobuloides 148
Newtonia 191

amphichroa 181, 184, 185, 186

brunneicauda 181, 184, 185,

186, 191

fanovanae 175, 188, 194, 197
Ninox

superciliaris 180
Noronhia 61, 65, 67, 73, 89, 102
Notoedres 139

Ochlandra

capitata 35
Ochrocarpos 61, 63, 77, 79, 83,
98-99

orthocladus 11, 83

tsaratananae 73, 83
Ocotea 60, 61, 63, 65, 66, 67, 69,

86, 100
Octolepis 11, 94, 104
Odontosoria

melleri 27, 33
Oedignatha 1 1 6
Olea 69, 89
Oleaceae 89, 95, 102

Oleandra

distenta 27
Oliganthes 71, 82
Oncostemum 45, 60, 61, 63, 65,
67, 69, 71, 73, 77, 79, 88-89,
101-102
Ophioglossum

palmatum 27

pendulum 27
Orchidaceae 89-91, 95, 102
Oreochromis

mossambicus 150
Oriolia

bemieri 181, 182, 195, 196, 197
Oryzorictes 219, 221, 228

hova 201, 202, 203, 209, 211,
212, 219, 221, 222, 225,
243

talpoides 211, 219, 243

tetradactylus 2 1 1
Oryzorictinae 206, 222, 223
Oryzoryctes

gracilis 2 1 3
Otus

rutilus 180, 184, 185, 186
Oxylabes 191

madagascariensis 140, 181, 184,
185, 187, 191

Pace i us 115, 116

angulatus 115, 116-117, 118,
119, 120

elevatus 115, 117

griswoldi 115, 116, 117-119,
120

madagascariensis 115, 116, 118,
120

mucronatus 115, 119

quadridentalus 115, 116

quinteri 115, 118, 120

scharffi 115, 116, 119
Pachylaelapidae 139, 140
Paederia 71, 92
Pallistrophoroides 139, 140
Pandanaceae 49, 51, 90, 95
Pandanus 46, 65, 66, 67, 69, 71,

73, 75, 77, 90, 159, 172
Panicum 11, 79, 90
Paracara

typus 149, 150
Paracontias

holomelas 161, 168

milloti 161, 168
Paraleucophenga 129, 131
Paramicrogale

occidentalis 206
Pararhadinea

albignaci 162

melanogaster 168
Parasitiformes 138-141
Paratilapia 149

bleekeri 149, 150

polleni 149-150
Paroedura

gracilis 161, 167, 170
Passeriformes 140

284

FIELDIANA: ZOOLOGY

Pauridiantha

paucinervis 103
Pay era 103

beondrokensis 103
Pellaea

angulosa 27
Peperomia 69, 73, 77, 79, 90, 102
Phaner

furcifer 265, 267, 271, 273
Phedina

borbonica 180, 185
Phelsuma

guttata 161, 167, 170

laticauda 161, 167

/mrata 161, 167, 170, 172

madagascariensis 161, 167, 170

pus///a 161, 167

quadriocellata 161, 167, 170,
172
Philepitta 191

castanea 180, 184, 185, 188,
191
Phthiraptera 137
Phyllanthus 75, 77, 79, 85, 100
Phyllarthron 69, 71, 82, 98
Phyllastrephus 182, 190
Phymatosorus

scolopendria 27, 34
Pj/ea 65, 69, 94
Piper 102

Piperaceae 90, 95, 102
Pittosporaceae 90, 95, 102
Pittosporum 61, 69, 71, 73, 90,
102

ambrense 102

humbertii 102

polyspermism 102

verticillatum 102
Pityrogramma

calomelanos 27, 33

humbertii 27, 33
Plagioscyphus 61, 65, 71, 93
Platycerium

madagascariense 27
Platypelis

barbouri 160, 166, 170

grant's 160, 166, 170

occultans 160, 166, 170

pollicaris 166

tsaratananaensis 160, 166, 170

tuberifera 160, 166, 170, 172
Pleopeltis 11, 91

macrocarpa 27
Plethodontohyla

bipunctata 160, 163, 166, 170

coudreaui 160, 163, 166

minuta 166

notosticta 160, 166, 170

ocellata 160, 166

serratopalpebrosa 160, 163,
166, 170
Ploceidae 140
P/ocews

nelicourvi 181, 184, 185, 187,
191
Pneumatopteris

remotipinna 27

subpennigera 27, 35
Poaceae 46, 47, 49, 51, 62, 63, 65,

75, 77, 78, 79, 90, 95, 102
Podocarpus 53
Polyalthia 97

emarginata 97

perrieri 65, 67, 80
Polyboroides

radiatus 183, 185
Polyscias 47, 55, 67, 69, 71, 72,

73, 75, 77, 81, 97
Potameia 63, 67, 69, 71, 86, 99

crassifolia 67, 86
Por/ios

scandens 61, 63, 65, 81
Pouridiantha

paucinervis 69, 92
Proctophyllodidae 140
Propithecus

diadema 266

candidus 265, 267, 268, 271,

273
diadema 271
Proteaceae 102
Protium 98

Protorhus61, 71, 80, 97
Pseudoacontias

angelorum 161, 162, 168, 172
Pseudobias

wardi 181, 194, 196, 197
Pseudocossyphus 182
Pseudocyclosorus

pulcher 27, 35
Pseudophegopteris

cruciata 27
Pseudoscorpionida 137, 139
Pseudoxyrhopus

heterurus 162, 168

microps 162, 168, 171

tritaneatus 162, 168, 171
Psorospermum 75, 83, 99
Psychotria 69, 71, 92, 103

ankafinensis 103

lokohensis 103
Pteridophyta 90-91, 95
Ptera

catoptera 27

cretica 27

elongatiloba
var. remotivenia 27

pseudolonchitis 27

woodwardioides 27, 35
Ptychadena

mascareniensis 160, 166
Pycnonotidae
Pyrostria 73, 92
Pyrro^j'a

rhodesiana 21, 27

Rallus

madagascariensis 183
Randia

pseudozosterops 181, 186, 191,
194, 196, 197
Ranidae 160, 166

Pa5 233

ra/r M s 231, 232, 242-245, 246,
252, 255, 256, 258, 259
Ravenala

madagascariensis 65, 67, 94,
159
Reptilia 161, 167, 170
Rhacophoridae 160, 167, 170
Rheocles 144

pellegrini 144
Rheocloides

pellegrini 144
Rhinonyssidae 140
Rhizophoraceae 91, 96, 102-103
Rhombophryne

testudo 166
Riparia

paludicola 183
Rochonia 98
Rodentia 139
Rothmannia 61, 92
Rubiaceae 49, 51, 60, 61, 62, 63,
65, 67, 69, 71, 73, 75, 77,91-
93, 96, 103-104
Pwe///a 97
Rumohra

adiantiformis 27

lokohoensis 27, 35
Rutaceae 65, 93, 96, 104

diversifolia 103

henriettae 27, 35
Saldinia 69, 92, 103
Sapindaceae 61, 67, 93, 96, 104
Sap/Mm 69, 71, 85
Sapotaceae 49, 51, 75, 77, 93, 96,

104
Sarcoptidae 139
Sarothrura

insularis 180, 183, 186, 188

torquata 181, 184, 185, 186
Scaptodrosophila 127, 131, 132
Scaptomyza 131, 132
Scatophaga

bispinosa 106
Schefflera 55, 69, 71, 73, 75, 77,
79, 81, 97

rw/a 183
Schismatoclada 69, 71, 73, 75, 77,
79,91,92, 103

farahimpensis 103
Schizaea

dichotoma 27

pectinata 27, 33
Scinicidae 161, 168, 171
Scleria 67, 69, 75, 79, 83
Scolopia 67, 69, 71, 77, 79, 85-

86, 100
Scrophulariaceae 104
Selaginella

fissidentoides

fa fissidentoides 27

INDEX

285

goudotana

var. goudana 28, 33

hildebrandtii 28, 35

pervillei 28

polymorpha 28

unilateralis 28
Specie 55, 77, 79, 82, 98
Setifer 228

setosas 204, 205-206, 209, 211,
220
Sicydiinae 151-153
Sicydium 151

acutipinnis 151

fasciatum 151

franouxi 151

lagocephalum 151

laticeps 151
Sicyopterus

fasciatus 151, 152, 153

franouxi 151, 152, 153

lagocephalus 151, 152-153

laticeps 151

pranouxi 152
Siphonaptera 137, 139

rhodantha 46, 53, 61, 64, 65,
69, 84, 99
Smilacaceae 93, 96
Smilax

kraussiana 61, 63, 71, 93
Sophophora 130
Soricidae 2 1 1
Sorindeia

madagascariensis 97
Sphaerostephanos

arbuscula 33
Sphenomeris

chinensis 28, 33
Stegana 131
Stenochlaena

tenuifolia 28, 61, 91
Sterculiaceae 93, 96, 104

flagellaris 28
Streblus 65, 88
Strelitzaceae 94, 96
Streptocarpus 100
Streplopelia 189, 190

picturata 180, 186, 189, 191
Strongvlodon 100
Stumpffia 160, 163, 166, 172

grand/.? 160, 166, 170

psologlossa 166

roseifemoralis 160, 166, 170

tetradactyla 166, 169

tridactyla 160, 163, 166, 169
5mch.v

murinus 2 1 1
Suregada 99

Sxmphonia 46, 53, 55, 63, 64, 65,
71. 75, 77, 79. 83, 99

microphxlla 75, 83, 99
Syzygium 63, 89. 102

Tabernaemontana 46, 62, 63, 68,
69, 80, 97

eusepala 97
Tachybaptus

pelzelnii 183

ruficollis 180
Tambourissa 53, 61, 69, 71, 73,
75, 88, 101

parvifolia 101

purpurea 101

thouvenotii 101
Tarenna 104
7Vc7an'a

gemmifera 28

humbertiana 28, 29, 33

magnifica 28, 35
Tenrec 228

ecaudatus 204, 206, 209, 211,
220
Tenrecidae 138-139, 201-229
Tenrecinae 205, 222, 243
Terpsiphone

mutata 181, 184, 185, 187
Thelypteridaceae 21
Thymeleaceae 94, 96, 104
Tiliaceae 94, 96, 104
Timaliidae 140
77na 104

striata 104
Trachelas 1 1 5

madagascariensis 1 1 5
Treculia 65, 88
7>eran 189, 190

australis 180, 182, 183, 189
Trichilia 61, 63, 88
Trichomanes 28, 32, 38

bipunctatum 28, 35

bonapartei 28

borbonicum 28

cupressoides 28

digitatum 28

erosum 28, 35

lenormandii 28

longilabiatum 28, 33

meifolium 28

montanum 28, 35

rigidum 28, 35

speciosum 28
Trilepisium

madagascariense 61, 63, 65, 8!
101
Trombiculidae 138-140
Trouessartiidae 140
Turdidae 140
Turnix

nigricollis 183

' eduardi 181, 187
Typhlopidae 162, 168
Typhlops

ocularis 162, 168
7>fo

soumagnei 188

f/apaca 46, 60, 61, 63, 66, 67, 71,

75, 85, 100
Ulmaceae 104
Uroplatus 161, 167, 172

efo-ziflH/ 161, 167, 170

fimbria tus 161, 167, 170

lineatus 161, 167

s/torae 161, 167, 170
Urticaceae 65, 94, 96, 104

Vaccinium 75, 77, 79, 84, 99
curvirostris 181, 184, 185, 187

rwbra 271, 273
Veprw61, 67, 69, 77, 93, 104
Verbenaceae 94, 96, 104
Vernonia 55, 67, 71, 73, 75, 77,

82, 98
Viscaceae 96, 104
V7scw/n 61, 71, 73, 75, 77, 87, 104
Vittaria

ensiformis 28

humblotii 28

scolopendrina 28

gymnocaudus 140, 219, 231,
232, 233, 242, 243, 244,
246, 251, 254, 255, 258,
259, 260

Weinmannia 55, 69, 71, 73, 75,
77, 78, 79, 83, 99
humbertiana 75, 83

Xenopirostris

polleni 183, 195, 196, 197
Xiphopteris 21. 28, 33

mysuroides 28

oosora
var. micropecten 28

sikkimensis 28
Xolalgidae 140
Xvlopia 61, 65, 67, 80, 97
Xyridaceae 94, 96, 104
Xyris 75, 79, 94, 104

Zanthoxylum

madagascariense 61, 63, 93
Zaprionus 127, 129, 131, 132

kolodkinae 132

mascariensis 132

spinipilus 132
Zingiberaceae 10, 35, 94, 96
Zo/705aw/5

aeneus 162, 168

madagascariensis 161, 168, 171

ra/z/^s 161, 162, 168

subunicolor 161, 168
Zoonavena

grandidieri 180
Zostero/

maderaspatana 181, 184, 185
Zygothrica 129

286

FIELDIANA: ZOOLOGY

3 0112 050509006