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Full text of "The birds of southeastern Madagascar"

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F1ELDIANA 



Zoology 

NEW SERIES, NO. 87 



The Birds of Southeastern Madagascar 



Steven M. Goodman 1 2 
Mark Pidgeon 2 
A. F. A. Hawkins 3 
Thomas S. Schulenberg 1 4 

department of Zoology 
Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, Illinois 60605-2496 USA 

2 World Wide Fund for Nature 
Aires Protegees, BP 738 
Antananarivo (101), Madagascar 

^BP 8511 

Antananarivo (101), Madagascar 

Conservation International 
2501 M Street, NW, Suite 200 
Washington, D.C. 20037 USA 



Accepted August 20, 1996 
Published November 26, 1997 
Publication 1487 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



© 1997 Field Museum of Natural History 

ISSN 0015-0754 

PRINTED IN THE UNITED STATES OF AMERICA 






Table of Contents 

Abstract 1 

Resume 1 

Introduction 2 

The Setting and Study Sites 3 

The Setting of Southeastern Madagascar ... 3 

Geology 3 

Climate 5 

Brief Review of Human Colonization and 

Occupation of the Region 6 

Habitat Types of Southeastern Madagascar ... 7 

Study Sites 16 

Methods and Terminology 19 

Mist-netting 19 

Census Surveys 20 

Habitat Classification 20 

Species Classification 20 

Collections and Sight Observations 20 

Condition of Reproductive Organs and 

Skull Ossification 21 

Systematic Order and Nomenclature 22 

Malagasy Bird Names 22 

Malagasy Locality Names 22 

Soft Part Colors 22 

Species Accounts 22 

Analysis and Discussion 90 

General Overview of the Regional Avi- 
fauna 90 

Relative Densities of Birds Based on 

Mist-netting 95 

Elevational Distribution of Birds 100 

Utilization of Sisal Plantations by Forest 

Birds 102 

Faunistics and Biogeography 105 

Conservation Problems in Southeastern 

Madagascar Ill 

Acknowledgments 116 

Literature Cited 117 

Appendix 1 . Gazetteer of Localities 

Mentioned in the Text 1 22 

Appendix 2. Names of Plant Genera, Spe- 
cies, and Families Mentioned in the 

Text 125 

Index to Malagasy Vernacular Bird 

Names 127 

Index to Scientific Names 128 



List of Illustrations 

1 . Map of southeastern Madagascar 4 

2. Cross section across southeastern Mad- 
agascar 4 



3. View from summit of Pic Trafonaomby 
(1956 m), with bank of clouds rolling 

in 7 

4. Littoral forest near Petriky, looking 

west 8 

5. Strand forest at the border between the 

sea and littoral forest at Manafiafy 9 

6. Lowland forest in the RNI 
d'Andohahela (parcel 1) at 400 m 10 

7. Lowland gallery forest in the RNI 
d'Andohahela (parcel 1) at 400 m 11 

8. Natural landslide in the RNI 
d'Andohahela (parcel 1) at 800 m 12 

9. Sclerophyllous forest on exposed ridge 
in the RNI d'Andohahela (parcel 1) at 
1700 m 13 

10. Mossy forest in the RNI d'Andohahela 
(parcel 1) at 1850 m 14 

1 1. Spiny forest in the RNI d'Andohahela 
(parcel 2) 15 

12. View across Mandrare River toward 
gallery forest on the east bank 16 

13. Secondary marsh habitat near Manafia- 
fy 17 

14. Pastureland converted from humid for- 
est at the northern boundary of parcel 

1 of the RNI d'Andohahela 18 

15. Young sisal plantation near Amboasary- 

Sud 103 

16. Total number of individuals per species 
recorded in sisal plantation along sur- 
vey transects 104 

17. Cluster analysis of faunal similarity of 
resident forest birds found at various 

sites on Madagascar 1 10 

1 8. Remaining portion of the Analalava 
Forest 113 

19. Charcoal pit in the Mandena Forest .... 115 



List of Tables 



1 . Observations of Milvus migrans on morn- 
ing transects in the Malaza Forest 29 

2. Observations of Coracopsis vasa and 
C. nigra on morning transects in the 
Malaza Forest 44 

3. Observations of Agapornis cana on 
morning and evening transects con- 
ducted in the Reserve Privee de Beren- 

ty 45 



in 



10. 



II 



External measurements (in mm) of 
Coua cristata maxima and Coua cris- 

tata pyropyga 48 

Number of Phyllastrephus spp. netted 

at humid forest sites 64 

Bird species recorded at each of the 

study sites 92 

Avifaunal composition of study sites in 

the RNI d'Andohahela 95 

Distribution of birds along an eleva- 
tional transect in the RNI 

d'Andohahela (parcel 1) 96 

Summary of mist-netting results at 

study sites 98 

Mist-netting summary in the littoral 

forests and spiny forest 99 

Comparison of bird contact frequencies 
in the littoral forests of Manafiafy and 
Itapera based on dawn censuses 100 



12. Mist-netting summary in the lowland 
humid forests 101 

13. Mist-netting summary of birds cap- 
tured along an elevational gradient in 

the RNI d'Andohahela (parcel 1) 102 

14. Sisal use by birds adjacent to the Mal- 

aza and Bealoka forests 104 

15. Distribution of resident forest birds at 
several well-known sites in humid for- 
est and spiny forest 106 

16. Faunal similarity indices using a bio- 
logical species concept of resident for- 
est-dwelling birds 109 

17. Faunal similarity indices using a phylo- 
genetic species concept of resident for- 
est-dwelling birds 109 

1 8. Comparison of bird-netting capture rate 
at relatively intact forested sites in the 
forests of Analalava and Marovony .... 114 



IV 



The Birds of Southeastern Madagascar 

Steven M. Goodman Mark Pidgeon 

A. F. A. Hawkins Thomas S. Schulenberg 



Abstract 

Southeastern Madagascar, defined here as the region from Tolagnaro north to Manantenina 
and west to the Mandrare River and its upper tributaries, contains a remarkable variety of 
habitats, including humid forests, dry spiny bush, littoral forests, coastal zones, high mountains, 
and areas of inland freshwater habitat. Within this region and its variety of habitats 189 bird 
species have been recorded. This represents 68% of the birds known to occur on Madagascar, 
within a region representing approximately 10,000 km 2 , or about 1.7% of the total land area 
of the island. 

Information is presented on the distribution, general aspects of natural history, diet, breeding, 
weight, soft part colors, and local names of the region's avifauna. This information is based 
on our own field work, published and unpublished observations, and museum specimens. 

The abrupt ecotone between wet and dry over a distance of a few kilometers is largely due 
to the north-south aligned Anosyenne Mountains, which act as a rain barrier or pluviometric 
fault. This shift in habitats over a short distance has few parallels elsewhere in the Old World 
tropics or subtropics and is reflected in extensive bird species turnover. 

Virtually all natural habitats within the region are currently threatened as a result of human 
activities. Little remains of the once extensive lowland forests on lateritic soils as a result of 
clearing for swidden agriculture, and the spiny forest has been extensively exploited for char- 
coal production and cleared for sisal plantations. The major reserve within the area is the 
Reserve Naturelle Integrate d'Andohahela, which is composed of three parts: parcel 1 is humid 
forest (63,100 ha), parcel 2 is spiny forest (12,420 ha), and parcel 3 is transitional forest (500 
ha). The future is bleak for natural habitats that remain outside the current protected areas 
system. 

Resume 

Le sud-est de Madagascar, defini ici comme la region comprise entre Tolagnaro au sud et 
Manantenina au nord et limited a 1'ouest par le fleuve Mandrare et ses affluents, abrite une 
remarquable vari&e" d'habitats naturels, depuis la foret pluviale sempervirente au bush £pineux 
sub-aride, en passant par la foret littorale, la zone cotiere, les hautes montagnes at les eaux 
douces continentales. 

Au sein de cette diversity de milieux naturels que pr£sente cette region, 189 especes 
d'oiseaux ont 6t6 repertories. Cela reprdsente 68% du total des especes d'oiseaux inventorizes 
a Madagascar. La region couvre approximativement 10,000 km 2 , soit environ 1,7% de la surface 
totale de Madagascar. Des informations relatives a la distribution, a 1'histoire naturelle, au 
regime alimentaire, a la reproduction, au poids, a la couleur des parties molles et aux noms 
vernaculaires malgaches de 1'avifaune de cette region sont apportdes. Les informations presen- 
tees sont issues de la synthese de travaux de terrain originaux, de donn£es scientifiques pubises, 
de donnetes scientifiques non-publtees et de donnetes mus£ologiques. 

La netted de I'ecotone constate sur seulement quelques kilometres entre les habitats humides 

FIELDIANA: ZOOLOGY, N.S., NO. 87, NOVEMBER 26, 1997, PP. 1-132 



at les habitats sub-arides est principalement le resultat de la presence de la chaine Anosyenne 
qui, de par son orientation nord-sud, fait office de barriere de pluie. Ce brutal changement 
d'habitat sur une courte distance occasionne un renouvellement important des especes 
d'oiseaux, phenomene constate au sein d'autres sites tropicaux et sub-tropicaux de l'Ancien 
Monde. 

Pratiquement tous les habitats naturels presents dans la region sont menaces de disparition 
de par les activites humaines. II ne subsiste que de petites surfaces de foret pluviale semper- 
virente de basse altitude sur sols lateritiques suite a la pratique de la culture itinerente sur brulis 
et le bush epineux sub-aride a vu sa superficie reduite par la production de charbon de bois et 
la culture du sisal. La principale aire protegee rencontree au sein de cette region est la Reserve 
Naturelle Integrate d'Andohahela composee de trois parcelles: la parcelle 1 est foret pluviale 
sempervirente (63,100 ha), la parcelle 2 est bush epineux sub-aride (12,420 ha), et la parcelle 
3 est foret de transition (500 ha). L'avenir des habitats naturels localises en dehors du systeme 
d' aires protegees est fortement hypotheque. 



Introduction 

The natural ecosystems of Madagascar contain 
a remarkable diversity of habitats, including large 
but diminishing expanses of lush tropical forests, 
high mountain alpine zones, and almost surreal- 
istic spiny bush. Reflected in this diversity of hab- 
itats is an avifauna that, although not as diverse 
as on other tropical islands (e.g., Borneo), shows 
a remarkably high level of endemism. Of the 204 
extant resident bird species known from the island 
(Langrand, 1990; Langrand & Appert, 1995; 
Goodman et al., 1996), 106 breed only on Mad- 
agascar and 25 also occur on neighboring islands 
(Comoros, Mauritius, and Reunion). Thus, about 
half of the avifauna is strictly endemic to Mada- 
gascar, and almost two-thirds (64%) is restricted 
to the greater Malagasy region. 

Much of the habitat diversity of Madagascar is 
compressed into the island's southeastern corner. 
Although much of this region is south of the Tro- 
pic of Capricorn, the humid forests are typically 
tropical in structure and species composition. The 
abrupt ecotone between wet and dry over a dis- 
tance of a few kilometers is largely due to the 
north-south-aligned Anosyenne Mountains, 
which act as a rain barrier or pluviometric fault 
(Battistini, 1964) for weather systems moving in 
from the Indian Ocean. Diminishing precipitation 
associated with this rain shadow has a dramatic 
effect on the floristic structure and composition 
across this zone. This shift in habitats over a short 
distance has few parallels elsewhere in the Old 
World tropics and is reflected in extensive bird 
species turnover. 

The southern limit of Madagascar's humid for- 
ests is reached on the windward side of the An- 



osyenne Mountains. Here one may be surrounded 
by 30-m-tall trees with large buttressed roots, the 
soils are rich in organic material, and terrestrial 
leeches (an indication of high humidity) are a 
common occurrence. The local avifauna is com- 
posed typically of humid forest species. From a 
few exposed ridges, on the leeward side of the 
Anosyennes, one can see to the immediate west, 
within a few kilometers, dry forest with its char- 
acteristic baobab (Adansonia) trees and thick 
stands of spiny Didiereaceae. From such vantage 
points one can hear humid forest birds calling in 
the immediate vicinity while the sounds of the dry 
forest emanate from below. This abrupt and dra- 
matic ecotone between wet and dry makes south- 
eastern Madagascar so fascinating and different 
from other areas of the island. 

No general synthesis on the birds of southeast- 
ern Madagascar exists. A. Grandidier visited the 
region, and various ornithological records were 
presented by Milne Edwards and Grandidier 
(1879). The Mission Zoologique Franco- Anglo- 
Americaine to Madagascar (1929-1931), which 
forms the basis for our modern working knowl- 
edge of the island's avifauna (Rand, 1936), did 
not visit this area. Over the past decade there has 
been an increase in ornithological activity on 
Madagascar, and numerous important records 
from the extreme southeast have been incorporat- 
ed in the works by Langrand (1990) and Langrand 
and Sinclair (1994). 

The purpose of this monograph is to summarize 
aspects of the natural history and distribution of 
bird species occurring in southeastern Madagas- 
car. We discuss numerous aspects of bird ecology 
and focus on documenting the remarkable species 
turnover across the pluviometric fault. 



FIELDIANA: ZOOLOGY 



The Setting and Study Sites 

The Setting of Southeastern Madagascar 

Within the limits of this study we define south- 
eastern Madagascar as the region from Tolagnaro 
north to Manantenina and west to the Mandrare 
River and its upper tributaries (Fig. 1). All refer- 
ences refer to this area unless otherwise stated. 

Geology 

The geology of southeastern Madagascar is 
complex, and the "Fort-Dauphin" group is one of 
the most intense examples of metamorphism and 
uplifting on the island (Brenon, 1972; Bazot, 
1974). The landscape of the region is dominated 
by two ranges, the Vohimena and Anosyenne 
mountains. The former starts just north of Tolag- 
naro. The eastern foot of the Vohimena chain runs 
north, parallel to the eastern sea coast but some 
2-8 km inland, to just south of Manantenina. The 
eastern foothills of the Vohimena Mountains rise 
out of the coastal plain and form an abrupt tran- 
sition from the sandy littoral zone to areas resting 
on lateritic soils. At several sites along this front 
the surface soil types change, typically with alti- 
tude, over a short ground distance. This shift in 
soils affects both the botanical and zoological 
communities. 

The mountains are formed from Precambrian 
gneiss and granitic rocks, and their deposited al- 
luvium is largely lateritic or ferrallitic soils (Bour- 
geat, 1972). The higher peaks of this range in- 
clude Pic Vohamena (1358 m) and Pic Vohimena 
(1 173 m). The eastern slopes of the range descend 
into numerous relatively small rivers that drain in 
a steep and short trajectory directly into the Indian 
Ocean. Along the eastern coast, at the base of the 
Vohimena Mountains, are a series of sediments 
dating from the Pleistocene, although these de- 
posits are often mixed with sands of various ages. 
The main forests of Mandena and Manafiafy rest 
on Karimbolian and Flandrian dunes (Battistini, 
1964). 

The Anosyenne Mountains are to the west of 
the Vohimena Mountains and run more or less 
parallel along a southwest-northeast axis from 
just west of Ranopiso to the Isandra Valley at 
the base of the Midongy-Sud Massif (Battistini, 
1964; Paulian et al., 1973). They have the same 
general geological history and composition as 
the Vohimena Mountains. The Anosyenne 



Mountains are distinctly higher with numerous 
summits over 1800 m (e.g., Pic Trafonaomby 
[1956 m] and Pic Andohahela [1935 m]). The 
eastern slopes of the range form a precipitation 
barrier to weather systems moving over Mada- 
gascar from the open sea. These mountains pro- 
vide the source for the Manampanihy and Efaho 
rivers. The former river drains towards the 
northeast and enters the sea at Manantenina, and 
the latter runs almost due south and meets the 
sea just west of Tolagnaro. Several small trib- 
utaries from the western slopes of the Vohimena 
Mountains also drain into the Manampanihy 
River. The western slopes of the Anosyenne 
Mountains form the source of the Mananara and 
Manambolo, which merge into the Mandrare 
River (Chaperon et al., 1993). This river, which 
enters the sea just south of Amboasary-Sud and 
270 km from its start near Pic Trafonaomby, is 
the lifeblood of thousands of people living in 
the arid zone to the west of the Anosyennes. 
Soils to the immediate west of the Anosyenne 
Mountains are typically lateritic clays and 
abruptly shift at the Androy sedimentary region 
to silicaceous sands. This region is geologically 
complex with the juxtaposition and infolding of 
numerous formations (Noizet, 1953). 

The main upper spines of the Vohimena and 
Anosyenne mountains are separated by a distance 
of less than 15-25 km either side of the Rano- 
mafana-Sud valley (Fig. 2). This valley is the con- 
duit of the Manampanihy River, and the soils are 
largely metasediments formed by erosion of sur- 
rounding mountain systems. The only remaining 
forested connection between the two mountain 
ranges is north of Isaka-Ivondro and south of the 
Ranomafana-Sud valley and is composed of a se- 
ries of ridges along a latitudinal axis consisting of 
the Col de Tsitongambarika, Col de Manangotry, 
and Col de Tanatana. 

West of the Anosyenne Mountains is a large 
well-drained basin of diminishing rainfall, low-ly- 
ing relief, and largely xerophilous vegetation. The 
basin covers an area of approximately 12,600 km 2 
and is predominantly drained by the Mandrare 
River. The basin has rather distinct geological 
boundaries delineated by the Anosyenne Moun- 
tains to the east, the extensive Manambian cliff 
cuestas of tectonic origin to the north, and the 
shallower escarpment leading to the Ambovombe 
pan just west of the Mandrare River. This river 
valley is the lowest portion of the basin, often 
bordered in areas by alluvial floodplains. There is 
a gradual increase in altitude from the coast to the 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 






KEY 




■ town or 


village 


A 


peak 




♦ 


forest 




HN 


Route N 


ationale 




Ifotaka TV\*. 




+ Itapera Forest 



Bealoka"!) Behara 



V 6 



[Ankapoky Forest k. J 



Amboasary 



♦ \ Manantantely Forest* 



"fcW Petriky* 



^ 



Ambovombe 



47 00 
I 



10 20 30 40 SO kl 
_l I I I I 



Fig. 1. Map of southeastern Madagascar. The reference to "parcel" designates the three parcels of the Reserve 
Naturelle Integrate d'Andohahela. 




Fig. 2. Cross section across southeastern Madagascar along a trajectory running from north of Manafiafy north- 
west to the Mandrare River. (Adapted from Roche & Marchal, 1955-1956.) 



FIELDIANA: ZOOLOGY 



Man a in hum Escarpment, 140 km to the north. 
Nevertheless, even at the base of these cliffs the 
altitude is slightly more than 300 m. The highest 
peaks in this region are the Vohimainty and Vohi- 
dagoro hills in parcel 2 of the Reserve Naturelle 
Integrate (RNI) d'Andohahela, which rise to 1005 
m. These hills are vestiges of the southern flank 
of the Anosyenne Mountains. In the north of the 
basin are the volcanic outcrops of the Vohidava 
ridge (922 m) and the Vohitsiombe mesa (904 m). 

The major geological influences affecting the 
region's relief include the Precambrian crystalline 
basement characterized by the highly metamor- 
phic Androyan system of volcanic intrusions 
mainly formed in the Upper Cretaceous and sec- 
ondarily in the Late Tertiary and Quaternary. 
There is also a sedimentary shelf of Permian to 
Recent origin that dominates the coastal region 
and extends into the lower Mandrare basin (Bat- 
tistini, 1972; Brenon, 1972). The relief in the 
northeast of the basin from Tsivory south to the 
confluence between the Mandrare and the Andra- 
tina rivers is dominated by the volcanic Androy 
Massif. Spectacular ledges, ridges, and mesas 
stand up from the crystalline beds of the Androy- 
an system. 

East of the basalt flows of the middle Mandrare 
basin is an area of extreme metamorphism with 
rich mineral deposits. This fractured and crystal- 
lized landscape is sometimes referred to as the 
Tranomaro group (Brenon, 1972), partly because 
the pediplains in the region of Tranomaro show 
typical consequences of the erosion of the Pre- 
cambrian shelf (Battistini, 1972). 

The lower Mandrare basin forms the eastern 
limit of the coastal sedimentary region. Much ev- 
idence is seen of recent sandstone and top sand 
deposition, particularly in the littoral zone and the 
lower continental shelf region of the western ba- 
sin. There is a gradual replacement in the south 
of lateritic clays, not only because they are sus- 
ceptible to accelerated erosion (Brenon, 1972; 
Jenkins, 1987), but also because they are overlain 
by more recent sedimentary deposition. 

The dominant features of the Mandrare Valley 
are the considerable deposits of red sands. These 
generally take on two forms: (1) red soils over- 
lying Cretaceous basalts and (2) red soils (colored 
by iron hydrates) that are essentially silicaceous 
sands (Jenkins, 1987). 

The littoral zone around the mouth of the Man- 
drare River, extending a few kilometers to the east 
and extensively to the west, is an area of substan- 
tial Quaternary dune accumulation. The beach at 



the ecotone between the sea and Lac Anony, for 
example, is a formation (still growing) of "liv- 
ing" white dunes referred to as the Flandrian re- 
gression (Battistini, 1972). To the west of Lac An- 
ony are deposits of alluvial sands brought down 
the Mandrare River, to the east of the lake are 
deposits of the rubified dune system of the Tat- 
simian Period (early Quaternary), and to the north 
of the lake are deposits of red paleosols formed 
during a pluvial period of the Neogene (Besairie, 
1970; Battistini, 1972). Besides the Tatsimian de- 
posits, which extend inland as far as Ifotaka, the 
coastal formation is largely comprised of Karim- 
bolian dunes of recent origin. These are the wide- 
spread dunes that extend around the southeastern 
coast north to Manafiafy. 

The vegetation is determined by the interaction 
of climate and substrate. Thus, for example, the 
spiny forest is characterized by such species as 
Alluaudia procera, A. ascendens, and Adansonia 
za on red or rubified soils, whereas the more 
coastal and well-drained Quaternary and Recent 
sands are typified by increasingly xeromorphic 
plants such as Euphorbia stenoclada, Alluaudia 
comosa, and Aloe vaotsanda. Gallery forest along 
the Mandrare and Mananara rivers generally 
grows on alluvial sands deposited by these rivers. 



Climate 

Rainfall — The most noticeable climatic fea- 
ture of southeastern Madagascar is the rapid de- 
crease in the amount of rainfall from east to west. 
To a much lesser extent there is a parallel decrease 
in precipitation on a north-south axis along the 
eastern coast. The moist easterly winds that hit 
the coast and Anosyenne Mountains provide or- 
ographic rainfall to the windward side of the 
mountains, whereas the leeward side to the west 
is in the rain shadow. Donque (1972, p. 136) sum- 
marized this transition zone: "the boundary be- 
tween the semi-arid climate and the tropical damp 
climate of the south-east coast is extremely sharp, 
along a 'pluviometric fault', which runs along the 
line of the Anosy range: over the distance of some 
sixty kilometres as the crow flies, there is a tran- 
sition from mean annual rainfalls of less than 600 
mm to amounts in excess of 1500 mm." For ex- 
ample, at Tolagnaro the annual rainfall is about 
1,500-1,800 mm; at Esira, 80 km to the northwest 
and west of the pluviometric fault, the annual 
rainfall is 740 mm; and at Behara, about 60 km 
west of Tolagnaro and in the heart of the south- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



eastern spiny bush, the annual rainfall is 530 mm 
(Paulian et al., 1973; Donque, 1975). The pluvi- 
ometric fault is much more abrupt than reflected 
above; the rate of meteorological change is ob- 
scured by the lack of weather stations in the tran- 
sition zone. Further, no data are available from the 
summit zone of the Anosyennes, which is very 
humid, with perhaps as much as several meters of 
rainfall per year. The Reserve Privee (RP) de Ber- 
enty, 70 km west of Tolagnaro and 10 km south- 
west of Behara, received approximately 491 mm 
of rainfall in 1984 and 426 mm in 11 months of 
1985 (Pidgeon, unpubl. data). RP de Berenty is 
the western limit of the area covered in this mono- 
graph. Perhaps the predictable aspect of the pre- 
cipitation on the southern spiny bush is its unpre- 
dictability. Long periods of negligible amounts of 
rainfall causing sustained drought, followed by 
significant amounts of rain that provide much of 
the year's total in a period of a few days, are com- 
mon. 

Along the north-south trajectory near the coast 
at Nahampoana (7 km north of Tolagnaro) the an- 
nual rainfall is 2, 1 30 mm, and at Manantenina (80 
km north of Tolagnaro) it is 3,000 mm (Paulian 
et al., 1973). Along this axis, precipitation prob- 
ably increases substantially inland and at higher 
altitudes. To a large extent the phytogeographical 
zones of the lowlands are directly correlated with 
rainfall pattern. 

The crest of the Anosyenne Mountains, partic- 
ularly along the eastern margins, is often shrouded 
in clouds (Fig. 3). This weather pattern is related 
to movements of humid air masses up from the 
east and the rapid descent of currents to the west 
into the hot and dry spiny bush (Humbert, 1935). 
These systems give rise to cooler temperatures, 
higher rainfall, and periods of thick fog in the 
summit areas. There also appear to be warmer air 
currents rising up the western slopes of these 
mountains. The cooler saturated air finds an ef- 
fective barrier that it infrequently crosses, and 
when it does cross, precipitation is often evapo- 
rated off into the atmosphere (Ratsivalaka-Ran- 
driamanga, 1985). 



Brief Review of Human Colonization and 
Occupation of the Region 

The first evidence of humans in southeastern 
Madagascar dates from the 9th century (Rako- 
toarisoa, 1997; Wright & Rakotoarisoa, 1997); 
this is about 800 years later than the earliest 



known human occupation of the island (MacPhee 
& Burney, 1991). Apparently, during the period 
from the 9th to the 12th century there were scat- 
tered small settlements along the coastal zone and 
along river valleys. Human subsistence relied pri- 
marily on fishing and cattle. There is no evidence 
that rice was grown during this period. The 13th 
and 14th centuries saw an increase in the size of 
villages and presumably a growth in the local 
population, as well as the presence of iron work- 
ing. Further, Chinese celadon ceramics found at 
several sites document trade contact with the out- 
side world. 

The period from the 15th to the early 17th cen- 
tury witnessed great cultural change in the region, 
particularly in the emergence of hierarchical so- 
cial organization (Wright et al., 1993). The ar- 
chaeological record indicates that communities, 
for example in the Efaho River Valley, were for- 
tified and there was a local influx of imported 
goods. This period is one of initial contact with 
Europeans, first the Portuguese, who established 
a fort in the region in 1540, then the French, and 
subsequently the Dutch (Decary, 1926). During 
this period there is evidence of irrigated rice, a 
practice that must have dramatically altered the 
freshwater wetlands of the region. Later in the 
17th century, guns were imported, there was in- 
creasing social complexity, and subsistence agri- 
culture was based on rice cultivation. 

Etienne de Flacourt, a representative of the 
French Compagnie des Indes Orientales in the lat- 
ter half of the 17th century who was based in 
Tolagnaro, was an excellent chronicler of cultural, 
social, and biological aspects of the region. Fla- 
court's treatise published in 1658 (reprinted edi- 
tion 1995) recounted in detail the effects of the 
political perturbations during this period on the 
local people of the region. He also described el- 
ephant birds and another animal interpreted to be 
a giant extinct lemur, suggesting that these species 
still existed in the region at that time or that at 
least memory of them lingered in local oral 
traditions. 

There is a rich modern oral cultural history 
from inland areas, including within and around 
the RNI d' Andohahela. A portion of this tradition 
probably dates from the 15th and 16th centuries 
(Charles, 1985; Razanabahiny, 1995). For exam- 
ple, the summit of Andohahela is reported to be 
the site where King Tehela sacrificed his son 
Mana, and even today this section of the reserve 
is considered taboo to enter. During our 1995 mis- 
sion to the eastern slopes of the RNI 



FIELDIANA: ZOOLOGY 




Fig. 3. View from summit of Pic Trafonaomby (1956 m) looking across the forested Anosyenne Mountains, with 
a bank of clouds rolling in from the east. Shrubs in the foreground are Philippia. (Photograph by N. Helme.) 



d'Andohahela, in areas without any evidence of 
preexisting trails or signs of recent human utili- 
zation of the forest, we found tombs. The often 
collapsed stone pillars forming these monuments 
were covered with thick layers of moss, and in 
several cases large trees were growing in the cen- 
ter of the tombs. Pottery found slightly below the 
ground surface at about 800 m was dated on sty- 
listic grounds to the 16th or 17th century (J. A. 
Rakotoarisoa, in litt.). On the basis of written ac- 
counts of voyagers who passed through the re- 
gion, the 17th century probably marks the era of 
substantial human habitation of the regional for- 
ests (Rakotoarisoa, 1997). Thus, our notion of ex- 
tensive untouched forest within the RNI 
d'Andohahela may be partly false. Rather, several 
hundred years of regeneration has been sufficient 
to hide the scars of previous human occupation. 

The 18th and 19th centuries were a period of 
considerable political change associated in part 
with the Merina (1825) and French (1896) colo- 
nizations of the area. These external powers dis- 
mantled the social and political structures of the 
local Anosy culture and imposed strict rule. In the 



20th century the ecological situation degenerated 
rapidly; regions described a few decades ago as 
forested are today no more than savannas. It has 
been estimated that 4,000 km 2 of the 7,000 km 2 
of the region was still forested at the beginning 
of this century (Rakotoarisoa, 1994), and although 
no accurate estimates are available, the current 
forest cover is substantially less than this figure. 
The Anosy economy remains largely agrarian 
(Peyrot, 1980), and swidden agriculture (tavy) re- 
mains a mainstay in the region. 



Habitat Types of Southeastern Madagascar 

Littoral Forest — Along the coastal margins 
of southeastern Madagascar, at less than 40 m 
above sea level, are a series of forests on sandy 
soils (Fig. 4) that comprise a distinct phytogeo- 
graphic unit (Ratsivalaka-Randriamanga, 1987; 
Lowry & Faber-Langendoen, 1991). This forest 
type is characterized by more than 1,000 mm of 
precipitation per year, a canopy height of between 
10 and 15 m, and a diameter at breast height (dbh) 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 




Fig. 4. Littoral forest near Petriky, looking west. (Photograph by T. S. Schulenberg.) 



of the largest emergent trees of generally less than 
50 cm. These littoral forests, which were presum- 
ably one of the major lowland forest types before 
the region was colonized by people, are presently 
confined to a few remnant parcels, for example 
near Bemangidy, Manafiafy, Itapera, Mandena, 
and Petriky. The latter forest parcel, 10 km west 
of Tolagnaro, is in a region with distinctly less 
rainfall than found in littoral forests on the east 
coast and subsequently has a different floral com- 
munity (Lowry & Faber-Langendoen, 1991). 

A subcommunity of the littoral forest, known 
as strand forest, is the vegetational zone imme- 
diately adjacent to the coastal beach. This forest 
type, which is often no more than 600 m wide, 
tends to be composed of halophytic plants. Pan- 
danus spp. and Casuarina are characteristic of 
this zone, particularly near Manafiafy, Itapera, and 
Mandena (Fig. 5). The general appearance of the 
littoral forest is of rather dense stands of short 
evergreen arborescents with stiff, rounded leaves, 
many of which are covered with a waxy cuticle. 
Wind desiccation and well-drained sandy soils 
may produce seasonal water stress. 

Humid Forest — The classification of this forest 



type is defined in various ways by botanists, and 
numerous alternative names are used. Humid for- 
ests, as defined here, are exclusively on lateritic 
soils in areas that receive at least 1,000 mm of 
rainfall per year, with a canopy height exceeding 
20 m tall, and with the largest canopy trees reach- 
ing 100 cm dbh. White (1983) divided this forest 
type into "rain forest" and "moist forest"; the 
former receives over 2,000 mm of rain per year 
and has a larger overall stature than the latter, 
which receives between 1,000 and 2,000 mm of 
rain per year. 

Humid forests can also be segregated into dif- 
ferent types based on elevation. The upper ele- 
vational limit of lowland humid forest generally 
occurs between 800 and 1000 m; several of our 
study sites (e.g., Marovony, Analalava, and Man- 
antantely) fall within this forest type. Forests 
above 600-800 m elevation are classified here as 
montane humid forest. With increasing elevation 
the stature of the forest decreases and the densities 
of bamboos, epiphytes, mosses, and tree ferns in- 
crease. The upper portions of montane humid for- 
est are often referred to as moss forest or upper 
montane humid forest. 



FIELDIANA: ZOOLOGY 




Fig. 5. Strand forest at the border between the sea and littoral forest at Manafiafy. The dominant tree is Pandanus. 
(Photograph by S. M. Goodman.) 



The following description of the plant com- 
munities and forest structure along an elevational 
transect conducted in parcel 1 of the RNI 
d'Andohahela in late 1995 is based on that of Hel- 
me and Rakotomalaza (in prep.). The forest at 400 
m had a canopy height of 15-25 m, and the dom- 
inant trees were Sorindeia madagascariensis, Ilex 
mitis, Syzygium, Oncostemon, Tambourissa spp., 
Dracaena reflexa, and various Rubiaceae (Fig. 6). 
Emergent trees, many of which had large but- 
tressed bases, reached 25-35 m, and the common 
species included Dilobeia thouarsii, Chrysophyl- 
lum boivinianum, Sloanea rhodantha, and Ocotea. 
Epiphytes were present, covering less than 20% 
of the available surface, and consisted of Asplen- 
ium, Pothos scandens, and various mosses. There 
was a high density of lianas. Large palms and 
bamboo clumps were not particularly common. 
Along riverbanks was a riparian plant community 
composed of Aphloia theiformes, Ficus, Antiro- 
hea, Weinmannia spp., Phyllanthus spp., and 
Dombeya spp. Ravenala madagascariensis was 
more common in light gaps along the river than 
in undisturbed forest (Fig. 7). 



At about 800 m there was a distinct floristic 
and structural change in the forest, most pro- 
nouncedly marked by an increase in the density 
and diversity of epiphytic plants. Canopy trees 
had at least 20-50% epiphytic cover, with Usnea 
lichens, Asplenium ferns, and Bulbophyllum or- 
chids dominating. Mosses were common. There 
also was a drop in the canopy height to between 
12 and 20 m and in the emergent trees to 20-25 
m. Canopy plants were dominated by Macaranga, 
Oncostemon, and the families Moraceae, Myrta- 
ceae, Clusiaceae, and Monimiaceae. Emergents 
included Sloanea rhodantha, Canarium obova- 
tum, Dilobeia thouarsii, Ocotea spp., and various 
Myrtaceae and Moraceae. The understory was 
dominated by Acanthaceae rather than Tambour- 
issa, as at 400 m. At 800 m, bamboo, particularly 
the lianescent Nastus, was common and formed 
dense tangles in light gaps. Large palms were 
rare, although small understory species were com- 
mon. In the area near our 800 m camp there was 
a high degree of community heterogeneity, sug- 
gesting strong environmental or edaphic gradi- 
ents. Further, there were clear signs of natural 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 







Fig. 6. Lowland forest in the Reserve Naturelle Integrate d'Andohahela (parcel 1) at 400 m. Note the large 
buttressed Sloanea and relatively light epiphytic growth on tree trunks and branches. (Photograph by M. Pidgeon.) 



landslides, the slopes of which had a different pi- 
oneering plant community than gallery or closed 
forest habitats (Fig. 8). 

Another major structural change occurred at 
about 1000 m, marking the shift from lowland to 
montane forest, which included a substantial in- 



crease in tree density, heavy epiphytic loads (50- 
80% cover), ground carpeted with spongy mosses, 
some areas of dense bamboo, and the near ab- 
sence of emergent trees. Canopy height varied 
from 12 to 20 m, and trees included Sloanea rho- 
dantha (with distinctly smaller root buttresses 



10 



FIELDIANA: ZOOLOGY 




Fig. 7. Lowland gallery forest in the Reserve Naturelle Integrate d'Andohahela (parcel 1) at 400 m. (Photograph 
by M. Pidgeon.) 



than at lower elevations), Canarium, Chrysophyl- 
lum boivinianum, and Croton monge. Lianas were 
still found in some areas but generally were less 
common than at lower elevations. The understory 
was relatively open and consisted of Acanthaceae, 
young saplings of canopy trees, Oncostemon spp., 
and in moister areas Cyathea spp. and Marattia 
fraxineae. 

At 1600 m the valleys contained moist montane 
forest and the ridges carried sclerophyllous forest 
(Fig. 9). Valley bottoms were dominated by Slo- 
anea rhodantha, often attaining a height of 30 m, 
and Strongylodon lianas; at the base of ridges 
Ravensara and Tambourissa were common; and 
on ridges in the sclerophyllous forest Dicoryphe 
viticoides, Tina isoneura, Elaeocarpus, Gaert- 
nera, and Aguaria were found. Moist areas were 
dominated by dense populations of Impatiens. 
Epiphytes were abundant, with 80-100% cover- 
age on horizontal branches. There were large 
clearings in the forest (0.5 ha), presumably caused 
by cyclone damage and landslides, and these areas 
were colonized by the pioneering tree species Ma- 
caranga and Dombeya. 



At 1900 m, on a plateau just below the summit 
of Trafonaomby, the forest was largely sclero- 
phyllous in nature with a high density of stems 
and low diversity, dominated by the families Ar- 
aliaceae, Lauraceae, Apocynaceae, and Flacour- 
tiaceae (Fig. 10). The understory was basically a 
monoculture of low-growing Acanthaceae with 
widely scattered patches of sedges. Epiphytic 
loads were heavy, approaching 100%. Lianas 
were rare, and palms were absent. The summit 
zone was covered with 3-m-tall sclerophyllous 
forest composed of Philippia spp., Alberta, Pit- 
tosporum, Aguaria, and Vaccinium. 

Transitional Forest — In the low foothills just 
west of the Anosyenne Mountains is a distinct 
vegetational structure referred to as transitional 
forest (O'Connor et al., 1985; Ratsivalaka-Ran- 
driamanga, 1987). One of the last remnants of this 
forest type is found in parcel 3 of the RNI 
d'Andohahela, the southern boundary of which 
borders Route Nationale 13 between Amboasary- 
Sud and Tolagnaro. In phytological characteristics 
this forest type is intermediate between humid and 
spiny forests. Mean annual precipitation in parcel 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



11 




Fig. 8. Natural landslide in the Reserve Naturelle Integrale d'Andohahela (parcel 1) at 800 m along the Andranohela 
River. Disturbed areas are colonized by Typha, Dombeya, Weinmannia, and Philippia. (Photograph by N. Helme.) 



3 is 700-800 mm per year (Nicoll & Langrand, 
1989), and the elevation varies between 100 and 
350 m (Eboroke, 1994). 

The vegetation of parcel 3 varies with edaphic, 
meteorological, and topographic conditions. In 
valley bottoms there is a multilayered forest with 



a 10- to 12-m-high canopy and relatively dense 
understory. The dominant plants are Millettia, 
Maba myriophylla, Commelina ramulosa, Ery- 
throxylum gerrardi, Cerbera venenifera, Dypsis 
decaryi, and Croton spp. (Eboroke, 1994; Drans- 
field & Beentje, 1995). On slopes canopy height 



12 



FIELDIANA: ZOOLOGY 




Fig. 9. Sclerophyllous forest on exposed ridge in the Reserve Naturelle Integrate d'Andohahela (parcel 1) at 1700 
m. (Photograph by N. Helme.) 



is about 10 m and the understory is dense, and 
the most common plants are Croton, Commelina 
ramulosa, Tarenna purinosum, Flacourtia luci- 
diaefolia, Diospyros myriophylla, Vepris sclero- 
phylla, and Alluaudia humbertii (Eboroke, 1994). 
The transitional forest of parcel 3 is probably the 
only reserve on Madagascar to have been desig- 
nated primarily for the protection of a single plant 
species, Dypsis decaryi, which is its most visible 
and distinctive aspect (Ratsirarson et al., 1996). 

Spiny Forest (= Sub arid Thorn Scrub) and 
Gallery Forest (= Riverine and Riparian For- 
est) — To the west of the Anosyenne Mountains 
and the transitional forest is a distinct forest type 
characterized by low precipitation (less than 700 
mm per year) and a fairly dense structure (Lowry 
& Faber-Langendoen, 1991). Along the Route Na- 
tionale 13 this forest type commences just west 
of the Col de Ranopiso, and slightly further west 
at Bevilany the flora is distinctly xerophytic (De- 
cary, 1927). Soils are usually lateritic, but at a few 
sites they are sandy. Our study site at Ankapoky 
is characteristic of this forest type, as is parcel 2 
of the RNI d'Andohahela (Fig. 11). The vegeta- 



tion of southeastern spiny forest is typically dom- 
inated by Alluaudia, Decaryia, Croton, Euphor- 
bia, Adansonia, Sarcostemma, Cynanchum, Ka- 
lanchoe, Pachypodium, Aloe, Delonix, Chadsia, 
Albizia, Crotalaria, Acacia, Commiphora, and, 
particularly on the rocky outcrops, Xerophyta. 
This is truly a deciduous forest. Leaf fall is ex- 
tensive in the spiny forest by the beginning of 
September, giving the appearance of an almost 
dead, petrified forest. Under conditions of severe 
water deprivation, some species will even drop 
branch segments. 

The spiny forest, often growing on higher and 
rockier ground with no permanent water table, re- 
lies on several adaptations to take advantage of a 
fleeting wet season with temporary groundwater. 
The ability of spiny forest plants to store water in 
tissues is key to their survival. Certain species 
have swollen trunks and thin leaves (e.g., Adan- 
sonia, Moringa, Pachypodium), or they have 
woody trunks and succulent leaves (e.g., Aloe, 
Kalanchoe). Some woody plants have extensive 
penetrating root systems and/or subterranean wa- 
ter storage vessels, and some succulent plants are 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



13 




Fig. 10. Mossy forest in the Reserve Naturelle Integrate d'Andohahela (parcel 1) at 1850 m and on plateau 
directly below Pic Trafonaomby. This largely sclerophyllous forest had a high density of stems and low plant species 
diversity, dominated by the families Araliaceae, Lauraceae, Apocynaceae, and Flacourtiaceae. The understory was 
dominated by low-growing Acanthaceae with occasional patches of sedges. (Photograph by N. Helme.) 



caulescent photosynthesizers with green stems 
(e.g., arborescent Euphorbia). Exceptionally, 
some plants (e.g., Alluaudia) exhibit succulent 
stems together with spines and succulent cadu- 
cous leaves. 

Throughout the spiny bush there are perma- 
nent and ephemeral river valleys, along the mar- 
gins of which grows gallery forest. The gallery 
forest largely is a woody forest that provides 
significant shade from canopy trees and taps a 
fluctuating water table. In contrast to the spiny 
forest, the gallery forest retains a semi- 
evergreen appearance all year. Important shade 
trees rarely drop all of their leaves, thus provid- 
ing protection for some more susceptible un- 
derstory plants. Some tree species also have the 
ability to fold their leaves (e.g., Tamarindus, 
Acacia, Bauhinia) during drought to minimize 
evapotranspiration. Gallery forest is dominated 
by a relatively narrow band of vegetation com- 
posed principally of Tamarindus indica, Acacia 
rovumae, Neotina isoneura, Celtis phillipensis, 



and Rinoria greveana. The Malaza Forest bor- 
dering the Mandrare River, at RP de Berenty, is 
typical of this forest type. Other dominant 
woody plants include Celtis gomphophyla, Al- 
bizia polyphylla, Crateva excelsa, Tabernae- 
montana, Ficus, and Bauhinia. The transition 
between gallery forest and adjacent spiny forest 
is usually abrupt (Fig. 12), although within the 
transition zone some species are shared. 

Freshwater — The lower slopes of the Ano- 
syenne and Vohimena mountains and the east- 
ern coastal plain contain freshwater lakes, 
streams, rivers, and marsh systems. The vege- 
tation of these aquatic systems varies consid- 
erably depending on soil type, water movement 
and depth, etc., but common elements include 
various grasses (Poaceae), sedges (Cyperaceae), 
water lilies (Nymphaea), pitcher plants (Nepen- 
thes madagascariensis), Pandanus spp., Rav- 
enala madagascariensis, and Typhonodorum 
(Lowry & Faber-Langendoen, 1991; pers. obs.) 



14 



FIELDIANA: ZOOLOGY 




Fig. 11. Spiny forest in parcel 2 of the Reserve Naturelle Integrate d'Andohahela. The dominant tree in the photo 
is Alluaudia ascendens (Didiereaceae). (Photograph by T. S. Schulenberg.) 



(Fig. 13). A considerable portion of freshwater 
habitat has been converted to rice paddy. 

Marine — The coastal beaches and lagoons pro- 
vide a wide range of habitats. Seabirds are regu- 
larly observed along coastal areas, particularly 
during and immediately after storms. Coastal 
sandy beaches and tidal estuaries provide habitat 



for resident and migrant shorebirds. Mangroves 
are found in a few areas, particularly along coastal 
lagoons and inland brackish rivers (Lowry & Fa- 
ber-Langendoen, 1991). 

Anthropogenic — By far the greatest portion 
of the land area of southeastern Madagascar 
represents not one of the natural habitats de- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



15 




Fig. 12. View across Mandrare River, looking east. On opposite bank is a narrow band of gallery forest. In the 
distance are the hills in parcel 2 of the Reserve Naturelle Integrate d'Andohahela. (Photograph by M. Pidgeon.) 



scribed above but rather some form of human- 
modified habitat, including grasslands, often 
used for cattle rangeland, formed by the cutting 
of forests and maintained by regular burning to 
prevent regeneration (Fig. 14); various types of 
secondary forest; exotic tree plantations, mostly 
Eucalyptus; and agricultural lands and garden 
plots ranging from extensive rice paddy areas to 
planted fields. Areas of forest cleared for swid- 
den agriculture are referred to as tavy. A wide 
variety of crops are grown in the region. 



Study Sites 

Field Work Conducted by M. Pidgeon 
(MP) — Between October 1983 and November 
1985 field studies were largely restricted to the 
gallery forests of Malaza and Bealoka with side 
trips to Bevala, Anjapolo, Ifotaka, and Lac An- 
ony. In October 1984 a brief survey commis- 
sioned by the World Wide Fund for Nature 
(WWF-US) was undertaken in the Bevilany and 
Fotsivolo Hills between Route Nationale 13 and 



the southern border of parcel 2 of the RNI 
d'Andohahela. During this period, some time was 
spent in the Andraraky Hills due east of Mokobe 
village. 

In 1984 a visit was made to parcel 2 of the RNI 
d'Andohahela in the area bordering Hazofotsy, 
and the following year this parcel was circumnav- 
igated on motorbike via Mokobe, Ankilitelo, and 
Ambatoabo. That same year the region along the 
Tanatana Trail (Isaka-Ivondro to Andonabe) and 
the Isedro Trail (Eminiminy to Mahamavo via Col 
d'Ambatomaniha) was visited. Between March 
1988 and September 1990 parcel 1 of the RNI 
d'Andohahela was explored more fully. Several 
exploratory trips were conducted, including the 
Tanatana Trail, the Isedro Trail to Evasia and Im- 
onty, the northern boundary trail between Enakara 
and Vohibaka, and the route from Esomony to 
Vohibaka (via Trafonaomby) and Marotsiva. 

In 1989 parcel 3 of the RNI d'Andohahela was 
circumnavigated and explored, with mammal trap- 
ping and mist-netting done in January 1990. In July 
and August 1989 a 3-day hike was undertaken to 
traverse parcel 2 of the RNI d'Andohahela from Ha- 



16 



FIELDIANA: ZOOLOGY 




Fig. 13. Secondary marsh habitat near Manafiafy. Ravenala madagascariensis (Strelitziaceae) is the banana-like 
tree in the center of the photograph. (Photograph by S. M. Goodman.) 



zofotsy to Ranomainty. This route passed via the 
summits of Vohidagaro and Vohimainty. In April 
1990, Tsivory (120 km north of Ambovombe) was 
visited to spot survey the foret classee, which in- 
corporates the Vohidava Hills. MP returned on the 



western side of the Mandrare River via Ebelo and 
then went on to Ifotaka. 
Field Work Conducted by S. M. Goodman 

(SMG) AND T. S. SCHULENBERG (TSS) IN 1989 AND 

1990 — Between September 1989 and early January 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



17 




Fig. 14. Pastureland converted from humid forest at the northern boundary of parcel 1 of the Reserve Naturelle 
Integrate d'Andohahela near Vohibaka. (Photograph by M. Pidgeon.) 



1990 (SMG and TSS) and between September and 
November 1990 (SMG), a zoological research group 
assessed the potential environmental effects of pro- 
posed mining of coastal mineral deposits in south- 
eastern Madagascar by QIT-Fer et Titane, Inc. 
(QIT). The primary role was to determine the spe- 
cies of birds occurring in this region; other members 
of the group conducted surveys of reptiles, amphib- 
ians, small mammals, and lemurs. In addition, other 
research teams performed similar surveys of the bot- 
any of the region. As a result of these studies, south- 
eastern Madagascar is biologically one of the best- 
known regions on the island. 

Study techniques are described below. The itin- 
erary at the main study sites, all in the Province de 
Toliara, Fivondronana de Tolagnaro, follows. 

Mandena Forest (littoral)— 8 km NE Tolag- 
naro, 24°58'S, 47°01'E, 20 m; 6-20 September 
1989— TSS. 

Petriky Forest (littoral)— 6.5 km SE Man- 
ambaro, 25°04'S, 46°53'E, 20 m; 22 September- 
2 October 1989— TSS. 

Manafiafy Forest (littoral)— 1.5 km NW 



Manafiafy (St. Luce), 24°47'S, 47°12'E, 20 m; 5- 
20 October 1989— SMG and TSS. 

Manafiafy Forest (strand forest) — 2.5 km 
SW Manafiafy (St. Luce), 24°48'S, 47°11'E, 40 
m; 21-25 October 1989— SMG and TSS; 18-22 
December 1989— SMG. 

Analalava Forest (humid) — 7 km N Manan- 
tenina, Foret d' Analalava, 24°13'S, 47°19'E, 40 
m; 28 and 29 October 1989 (reconnaissance) — 
SMG and TSS; 5-11 November 1989— SMG and 
TSS; 23-25 November 1990— SMG. 

Marosohy Forest (humid) — 3 1 October and 1 
November 1989 (reconnaissance) — SMG and 
TSS. Foret de Marosohy, along tributary of Tsi- 
tongatona River, along Enakara-Antseva forest 
trail, 15.5 km (by air) WNW Ranomafana-Sud, 
24°34'S, 46°48'E, 725 m; 23 November to 4 De- 
cember 1989 — SMG. Foret de Marosohy, along 
tributary of Tsitongatona River, along Enakara- 
Antseva forest trail, 15 km (by air) WNW Ran- 
omafana-Sud, 24°34'S, 46°49'E, 425 m; 4-14 De- 
cember 1989— SMG. 

Bezavona Forest (humid) — 1.5 km (by air) 



18 



FIELDIANA: ZOOLOGY 



NW Nahampoana, 7.5 km (by air) NNW Tolag- 
naro, Foret de Bezavona, 24°58'S, 46°58'E, 75 m; 
24 December 1989 (reconnaissance); 26-30 De- 
cember 1989; 17, 23, 25 September, 13 and 21 
October 1990 (day trips from Tolagnaro) — SMG. 

Manantantely Forest (humid) — 12.2 km (by 
air) NE Manambaro, 8.5 km (by air) NW Tolag- 
naro, Foret de Cascade, 24°59'S, 46°56'E. 100 m; 
15 September 1990 (reconnaissance) — SMG; 27 
September to 3 October 1990— SMG. 

\ n kapok\ Forest (spiny) — 13 km (by air) NE 
Amboasary-Sud, 21 km (by air) NW Ranopiso, 
Foret d'Ankapoky, 24°59'S, 46°31'E, 70-120 m; 
8-13 October 1990— SMG. 

Itapera Forest (littoral) — 19.5 km (by air) NE 
Tolagnaro, Foret dTtapera, 24°52'S, 47°07'E, 0- 
20 m; 15 September 1990 (reconnaissance) — 
SMG; 15-20 October 1990— SMG. 

Marovony Forest (humid) — 19 km (by air) 
NNE Manantenina, Foret de Marovony, 24°06'S, 
47°22'E, 50 m; 27 October to 4 November 
1990— SMG. 

Account of Field Work Conducted by SMG, 
E Hawkins (FH), and MP in Late 1995 — From 
19 October to 14 December 1995 a multidisci- 
plinary and multinational group of biologists con- 
ducted an inventory of the RNI d'Andohahela. 
The main focus of the project was to survey an 
elevational transect of the humid forest zone of 
parcel 1. Five sites, centered on camps at 440, 
810, 1200, 1500, and 1875 m, were the focal 
points of this mission. Further, a sixth site was 
surveyed in the spiny forest habitat of parcel 2. 

During this survey FH was the principal orni- 
thologist, and he conducted point counts (see Cen- 
sus Surveys, p. 20) and made general observa- 
tions. SMG and MP worked primarily with mam- 
mals and invertebrates but also had time for or- 
nithological observations, which were reported to 
FH. Mamy Ravokatra was responsible for the bird 
netting, and the captured birds were processed by 
SMG. In general we tried to use several different 
techniques to maximize data gathered and to ap- 
proach near completeness of the transect bird lists 
(Bierregaard, 1990; Remsen, 1994). 

Camp sites, all within the Province de Toliara, 
and inclusive dates of occupancy follow. 

Camp 1—19-28 October 1995. RNI 
d'Andohahela, parcel 1, 8 km NW Eminiminy, 
24°37.6'S, 46°45.9'E, 440 m. 

Camp 2 — 28 October to 7 November 1995. 
RNI d'Andohahela, parcel 1, 12.5 km NW Emi- 
niminy, 24°35.6'S, 46°44.3'E, 810 m. 

Camp 3—7-17 November 1995. RNI 



d'Andohahela, parcel 1, 13.5 km NW Eminiminy, 
24°35.0'S, 46°44.1'E, 1200 m. 

Camp 4—17-27 November 1995. RNI 
d'Andohahela, parcel 1, 15.0 km NW Eminiminy, 
24°34.2'S, 46°43.9'E, 1500 m. 

Camp 5 — 27 November to 5 December 1995. 
RNI d'Andohahela, parcel 1, 20.0 km SE Andra- 
nondambo, 24°33.7'S, 46°43.3'E, 1875 m. 

Camp 6—7-15 December 1995. RNI 
d'Andohahela, parcel 2, 7.5 km ENE Hazofotsy, 
24°49.0'S, 46°36.6'E, 120 m. 



Methods and Terminology 
Mist-netting 

At each site a series of 12-m mist nets was 
erected. The bottom edge of each net was within 
20 cm of the ground. The nets were monitored at 
regular intervals between sunrise and 1 hour after 
sunset. Nets were left open for 24 hours per day 
to sample nocturnal birds and bats. Mist-netting 
results are expressed as the number of individuals 
(or species) captured (NCI) per "net-day," de- 
fined as the continuous use of a 12-m net for a 
complete 24-hour period. 

Mist-netting provides a quantitative, although 
biased, estimate of bird relative abundance for 
species that are regularly active in the understory 
and lower middle story of the forest. The tech- 
nique imperfectly samples species that are too 
large to be restrained by the net and those that 
are primarily active in the middle or upper forest 
canopy. Further, it may misrepresent actual mea- 
sures of relative abundance for some species in 
the understory, depending upon their average 
flight distance and social system (Remsen & Par- 
ker, 1983; Remsen & Good, 1996). However, 
given standardization of technique and condi- 
tions, as well as exclusion of canopy species, 
mist-netting data can be useful to measure and 
compare understory species richness at different 
sites (Poulsen, 1994). 

All birds captured were brought back to the 
camp in cloth bags, weighed, and measured. A 
portion of individuals captured were prepared as 
study skins or anatomical specimens (skeletons 
or fluid preserved). Those individuals released 
either were banded with a plastic ring stamped 
with a unique number or a color sequence or 
were marked on the feathers with a permanent 
ink pen. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



19 



Census Surveys 



Habitat Classification 



Mist-netting data were supplemented with di- 
rect observations and auditory identification. Strip 
censuses were conducted at dawn at Mandena, Pe- 
triky, Manafiafy, and Itapera, and results present- 
ed are the number of individuals per species per 
unit length of trail. Such censuses allow for an 
estimate of relative abundance of bird species, 
particularly during the breeding season (approxi- 
mately September to December), when birds are 
vocalizing. These censuses were conducted at or 
immediately after dawn until approximately 
09h00, the period of maximum vocal activity for 
many bird species. At the other sites, detailed 
notes were kept on the birds observed or heard. 

During the 1995 expedition to the RNI 
d'Andohahela data were collected within an alti- 
tudinal band ± 100 m and less than 3 km in hor- 
izontal distance from camps, which were estab- 
lished at 440, 810, 1200, 1500, and 1875 m in 
humid forest in parcel 1 and in spiny forest in 
parcel 2. Species lists were compiled by direct 
observation while investigations walked along 
forest trails, by call-playback of bird calls using a 
tape recorder, by static observation from broken- 
canopy watch points, and by mist-netting. Obser- 
vations from canopy watch points (principally for 
raptors) were made between 07h00 and 1 1 hOO. No 
suitable site was found for canopy watches at 440 
m, but in other elevational zones at least 6 hours 
of observations were accrued per site. Point 
counts were made at 150-m intervals (in humid 
forest) or 200-m intervals (in spiny forest) along 
marked and measured forest trails. A minimum of 
12 point count sites were used within each tran- 
sect zone. Where possible, 5 point count sites 
were established at each site in each of the fol- 
lowing situations: ridge, slope, and valley bottom. 

Each point count site was sampled twice, once 
between 04h30 and 06h00 and once between 
06h30 and 09h00, but never on the same morning. 
During each sample count, which lasted for 10 
minutes, the following data on each bird contact 
were noted: species, estimated distance from ob- 
server (to nearest 10 m), nature of contact (song, 
call, wing noise, or visual), and time of contact. 
Densities were calculated using distance estimates 
for those species for which sufficient data were 
collected. Estimates of bird densities from the 
RNI d'Andohahela will be presented in a forth- 
coming monograph on the 1995 inventory of the 
reserve. 



For the analysis of bird habitat preference, we 
used the following broad categories: 

Forest (F) — Species restricted to continuous 
tracts of disturbed or intact forest. 

Open (O) — Species that are found in open ar- 
eas, including natural openings and tavy. 

Mixed (M) — Species that use forest and open 
habitats. 

Aquatic (A) — Species that are found in wet- 
land habitats, including the littoral zone and sea. 



Species Classification 

For the designation of a species' geographic 
distribution the following symbols have been 
used: 

* = endemic — species only found on Mada- 
gascar. 

(*) = endemic to region — species that are re- 
stricted to Madagascar, Comoros, and 
Mascarenes. 

N = nesting — species that breed on Madagas- 
car but are not endemic to the island. 

M = migrant — species breeding elsewhere but 
spending the austral summer on Madagas- 
car. 

I = introduced — species not native to Mada- 
gascar. 



Collections and Sight Observations 

A considerable amount of previously unpub- 
lished information is available in museum collec- 
tions on the birds of southeastern Madagascar. We 
have examined material in numerous institutions, 
and these data have been incorporated herein. 

amnh — American Museum of Natural History, 
New York. 

bmnh — The Natural History Museum, formerly 
British Museum (Natural History), Tring, United 
Kingdom. 

fmnh — Field Museum of Natural History, Chi- 
cago. 

mnhn — Museum National d'Histoire Naturelle, 
Paris. 

nhb — Naturhistorisches Museum, Basel. 

smf — Forschungsinstitut und Naturmuseum 
Senckenberg, Frankfurt am Main, Germany. 

pbzt — Pare Botanique et Zoologique de Tsim- 



20 



FIELDIANA: ZOOLOGY 



bazaza (at least in part a portion of the former 
ORSTOM collection), Antananarivo, Madagascar. 

From the time of earliest European colonization, 
Tolagnaro (Fort-Dauphin) has been an important 
trading port, and some of the earliest zoological 
collections to be exported from Madagascar came 
from there. For example, the first known bird col- 
lections made in the Tolagnaro region were by 
Pierre Poivre in 1756 (Stresemann, 1952). With 
such early collections, which are usually labeled 
"Fort-Dauphin," it is not clear if the specimens 
were actually collected in the immediate surround- 
ings of Tolagnaro or across a broader geographic 
zone, Tolagnaro merely being the port of exporta- 
tion. For the sake of simplicity we have cited such 
material as being from "Fort-Dauphin." 

As in so many areas of the island, the initial 
exploration of remote forested regions of south- 
eastern Madagascar was conducted by the French 
botanist Henri Humbert, who visited the region in 
1928 and in 1933-1934. On the basis of his dis- 
covery of intact forest in the mountain zones, part 
of the Andohahela Forest was designated as a re- 
serve (Humbert, 1941). Subsequently, the size and 
habitat types represented within the reserve were 
greatly expanded (Nicoll & Langrand, 1989). 

In 1931 Hans Bluntschli visited areas near Am- 
boasary-Sud and near Eminiminy, which was later 
gazetted as parcel 1 of RNI d' Andohahela 
(Bluntschli, 1932, 1933). The Bluntschli collec- 
tions were dispersed to (at least) amnh, nhb, and 
SMF (Bluntschli [1951]). The material housed in 
smf has been partially described by Dee (1986). 

In December 1948 Harry Hoogstraal visited 
southeastern Madagascar to collect material on 
vertebrate ectoparasites and to study their impor- 
tance as disease vectors (Hoogstraal, 1953; Uilen- 
berg et al., 1979). The main focus of his work 
was mammals, but some bird specimens were col- 
lected and sent to fmnh. Hoogstraal visited forests 
near Tolagnaro, Mandena, and Bemangidy. 

In 1971 and 1972 a group of scientists associ- 
ated with the Centre National de la Recherche 
Scientifique studied the ecosystems of the Ano- 
syenne Mountains (Paulian et al., 1973). Their re- 
port focused principally on the geomorphology, 
climate, and floristic structure. 

Other small research collections made in south- 
eastern Madagascar include those of John G. Wil- 
liams near Tolagnaro (National Museums of Ke- 
nya) and Georges Randrianasolo near Lac Anony 
and the interior portions of the spiny forest (pbzt). 
In 1948 Philippe Milon collected birds near To- 



lagnaro, Isaka-Ivondro, and at a site "30 km 
NNW Fort Dauphin," which was in or at the edge 
of parcel 1 of the RNI d' Andohahela (mnhn). In 
1977 Appert (1985) visited the Bemangidy Forest. 
Specimens taken in 1989 and 1990 during the QIT 
field work are at fmnh, and some have been 
transferred to the Service de Paleontologie and 
Departement de Biologie Animale, University 
d'Antananarivo. 

Rakotondranony's (1977) thesis on the verte- 
brate fauna of the RP de Berenty, particularly birds, 
contains numerous records of species previously 
not reported from southeastern Madagascar (e.g., 
Anas bernieri and Philepitta schlegeli) or highly 
exceptional in the Mandrare River basin (e.g., 
Alectroenas madagascariensis and Pseudobias 
wardi). Because no precise details were presented 
with his observations and consequently they are 
not verifiable, we have generally not included these 
doubtful records within the species accounts. 

We also gathered detailed, verifiable, but un- 
published information on birds observed by or- 
nithologists and bird-watchers that have visited 
the region. Observers are identified in the text by 
their initials: 

DT = David Thorns 

DW = David Waugh 

DWo = David Wolf 

FO = Frank Oatman 

GR = Georges Randrianasolo 

IS = Ian Sinclair 

LW = Lucienne Wilme" 

OL = Olivier Langrand 

SJ = Stig Jensen 

SOC = Sheila O'Connor 



Condition of Reproductive Organs and Skull 
Ossification 

Within the species accounts we have often in- 
cluded information on the reproductive state of 
collected birds based on the condition and devel- 
opment of the sex organs, and age was estimated 
from skull ossification. This information was used 
to infer aspects of breeding seasonality, age at first 
reproduction, and plumage sequences. However, 
as a caveat, we emphasize that there is not always 
a clear relationship between size of testes and 
sperm production, particularly in tropical birds 
(Moreau, 1936; Snow & Snow, 1964; Foster, 
1975). Further, there are also cases of passerines 
breeding at an unusually early age (often under 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



21 



extreme environmental conditions) and presum- 
ably before complete ossification of the skull 
(Gibbs et al., 1984). 



Marovony — Antaisaka, with some Tanosy in- 
fluence. 

RP de Berenty (= Malaza Forest) — Tandroy. 



Systematic Order and Nomenclature 

We generally follow the systematic order, no- 
menclature, and English vernacular names used 
by Langrand (1990). The Procellariiformes follow 
Jouanin and Mougin (1979). In a few cases we 
have deviated from this system. Subspecific de- 
terminations are based on our own comparisons 
of material taken in southeastern Madagascar. In 
cases when no material was available or patterns 
of geographic variation are not clear, the subspe- 
cific name is presented in brackets. For the sci- 
entific names and families of plants (Appendix 2), 
we have followed Missouri Botanical Garden 
(1993) and Mabberley (1989). 



Malagasy Bird Names 

We collected information on the bird names 
used by local people. In most cases this was done 
with the bird in the hand or clearly visible to the 
informant. Some of the difficulties in transcribing 
names from Malagasy have been reviewed by 
Langrand (1990, pp. 53-55). In southeastern 
Madagascar these problems are further exacerbat- 
ed by the fact that several cultural groups inhabit 
the region, often each with a variant of the pro- 
nunciation or even a totally different name for a 
given species. An index to local names is pre- 
sented at the back of this monograph. 

In many cases dialectal differences between 
these groups are not discrete, and rather than pre- 
senting names associated with various cultures we 
present them by locality. The following is a gen- 
eral guide to the cultural group/dialect associated 
with the various sites from which bird names are 
presented. 

RNI d'Andohahela (parcel 1) — Tanosy. 

RNI d'Andohahela (parcel 2) — Tanosy, Tan- 
droy. 

Bealoka — Tandroy. 

Hazofotsy — Tandroy. 

Manafiafy — Tanosy. 

Manantantely — Tanosy. 

Manombo — Antaisaka. 

Marosohy — mostly Tanosy, with some Antai- 
saka influence. 



Malagasy Locality Names 

Both French and Malagasy names currently are 
used for localities in Madagascar. In the vast ma- 
jority of cases we use the Malagasy name and 
spellings indicated on Foiben-Taosarintanin'i 
Madagasikara maps (FTM, Institut National de 
Geodesie et Cartographie). All locations men- 
tioned in the text are listed in Appendix 1 with 
respective latitude, longitude, and elevation. We 
have attempted to include all relevant locality 
synonyms. 

For the sake of brevity we have used abbrevi- 
ations for the category names of protected areas 
and several Malagasy governmental organiza- 
tions. 

DEF — Direction des Eaux et Forets. 

JIRAMA — Jiro sy Rano Malagasy. 

PN — Pare National. 

RNI — Reserve Naturelle Integrate. 

RP — Reserve Privee. 

RS — Reserve Speciale. 



Soft Part Colors 

Langrand (1990) summarized information on 
the soft part coloration of Malagasy birds. During 
our field work in southeastern Madagascar we 
have been able to supplement this information. 
Here we provide further details on the soft part 
colors of birds, based on live or freshly killed 
birds in the hand. Some information is presented 
on soft part colors inscribed on data tags of 
Bluntschli specimens. 



Species Accounts 

Procellariiformes 

Diomedeidae 

Diomedea cauta 

Shy or White-capped Albatross 

A single bird was observed from a promontory, 
just east of Tolagnaro, on 22 August 1 990 (Lan- 
grand & Sinclair, 1994). 



22 



FIELDIANA: ZOOLOGY 



Diomedea chlororhynchos 
Yellow-nosed Albatross 

On 16 July 1991, three Yellow-nosed Albatross 
were observed 1 5-20 km off the coast of Tolag- 
naro (IS; Langrand & Sinclair, 1994). 



Procellariidae 



Bulweha fallax 
Jouanin's Petrel 

On 22 December 1991, two Jouanin's Petrels 
were observed from a ship at sea northeast of To- 
lagnaro. The only other record of this species 
from the region was one individual on 12 January 
1992, 130 km northeast of Tolagnaro (Langrand 
& Sinclair, 1994). 



Daption capense 
Cape Petrel 

Two Cape Petrels were observed from a ship 
on 16 July 1991, 15-20 km off the coast of To- 
lagnaro and in view of land (IS; Langrand & Sin- 
clair, 1994). 



Pterodroma macroptera 
Great-winged Petrel 

A group of eight Great-winged Petrels was ob- 
served on 22 August 1 990 from a promontory due 
east of Tolagnaro, and another flock of six was 
noted on 22 December 1991 from a boat in view 
of land northeast of Tolagnaro (Langrand & Sin- 
clair, 1994). 



Puffinus pacificus 

Wedge-tailed Shearwater 

On 2 January 1990, during a cyclone, two 
Wedge-tailed Shearwaters were observed about 
300 m inland at Tolagnaro, and on 20 October a 
flock of 30 was observed just offshore (SJ). This 
species has been observed numerous times off the 
coast of Tolagnaro (Milon et al., 1973; Langrand, 
1990). 



Puffinus atrodorsalis 
Mascarene Shearwater 

The only record of this recently described spe- 
cies in southeastern Madagascar was a group of 
five observed 190 km NE of Tolagnaro on 28 Au- 
gust 1991 (Shirihai et al., 1995). 



Pterodroma mollis 
Soft-plumaged Petrel 

A group of four individuals was observed on 
16 July 1991 from a boat within sight of land and 
15-20 km off the coast of Tolagnaro (IS; Lan- 
grand & Sinclair, 1994). 



Pterodroma baraui 
Barau's Petrel 

One Barau's Petrel was noted on 22 December 
1991 off the coast of Tolagnaro within sight of 
land (IS; Langrand & Sinclair, 1994). 



[Pachyptila vittata 
Broad-billed Prion 

A group of over 30 prions was noted on 1 6 July 
1991 about 15-20 km off the coast of Tolagnaro 
(IS). They were most likely this species.] 



Podicipediformes 

Podicipedidae 

Tachybaptus pelzelnii 
Madagascar Little Grebe 

The Madagascar Little Grebe was rare in south- 
eastern Madagascar. On 7 August 1982 about 20 
individuals, all in nonbreeding plumage, were ob- 
served on a small pond between Lac Anony and 
Lac Erombo (OL & LW). This is the only known 
record from the region. 



Pelecaniformes 

Phaethonidae 

Phaethon aethereus 
Red-billed Tropicbird 

Milne Edwards and Grandidier (1879) reported 
that a Red-billed Tropicbird was collected at Tol- 






GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



23 






agnaro. The fate of the specimen is unknown to 
us. 



Phalacrocoracidae 

Phalacrocorax africanus pictilis 
Reed Cormorant 

The Reed Cormorant has a restricted range in 
southeastern Madagascar. This species has been 
observed along the Mananara River, near Hazo- 
fotsy, and on the Mandrare River during different 
times of the year, although it was not common 
and is not known to breed in this region. 
Bluntschli collected two individuals near Am- 
boasary-Sud on 22 October (amnh, smf). Up to 
seven individuals, including two juveniles, were 
observed roosting with Bubulcus ibis in trees 
along the Mananara River in parcel 2 of the RNI 
d'Andohahela in mid-December. 

Local Name — Razamboay (Berenty). 



Ciconiiformes 

Ardeidae 

Ixobrychus minutus subsp. 
Little Bittern 

The only report of a Little Bittern from the area 
was a male at Mandena on 20 September. This 
record could be of nominate minutus, a Eurasian 
migrant to sub-Saharan Africa, or of podiceps, the 
resident subspecies on Madagascar. 



Nycticorax nycticorax [nycticorax] 
Night Heron 

The Night Heron was noted along coastal river 
margins, lagoons, and marshes near Manafiafy, 
Itapera, Mandena, and Petriky. It was generally 
observed at dusk and dawn. A regularly used 
roost site was on the east bank of the Mandrare 
River, 4 km south of Amboasary-Sud, at Bevala. 
Up to two birds were observed at dusk flying 
along the Mananara River. 

Breeding — Night Herons nested in the RP de 
Berenty; at least 10 nests were occupied in early 
September 1984 and late August 1985. Juveniles 
were observed on 1 March along the Mandrare 



River near the RP de Berenty (OL) and on 20 
September near Bevala. 

Local Name — Fitatry (Manombo). 



Ardeola ralloides 
Squacco Heron 

The Squacco Heron was relatively common in 
coastal areas. We observed it at Itapera, Mandena, 
and Petriky. It also was occasionally noted in 
flooded fields, particularly rice paddies, in the 
Tolagnaro area, near the RP de Berenty, and 
Bealoka. 



Ardeola idae 

Malagasy Pond Heron 

This species was uncommon in agricultural ar- 
eas, particularly flooded rice paddy, in the Tolag- 
naro region. It also was noted along the Mananara 
River near Hazofotsy, in rice paddies near Am- 
boasary-Sud, and in open areas at the edge of the 
Marosohy Forest. We have no observation of this 
species in the region above 350 m. 



Bubulcus ibis [ibis] 
Cattle Egret 

The most common open country ardeid, the 
Cattle Egret was regularly noted foraging in ag- 
ricultural areas, particularly rice paddies, in pas- 
tureland, often with cattle, in flooded fields, and 
at the edge of marshes. It was more common in 
lowland areas below 80 m. Large numbers would 
search gregariously for insects, particularly or- 
thopterans, on the banks of the Mandrare River. 
Roosts of up to 30 individuals, including birds in 
breeding plumage, were noted along the Manan- 
ara River in mid-December. 

Breeding — Colonies were found in the RP de 
Berenty and along the Mandrare River. Apparent- 
ly this species has an irregular breeding season in 
this region. On 1 March a colony of about 100 
nests, with eggs and young, was found in the trees 
along the Mandrare River. On 23 December a 
nearby colony contained about 1,000 active nests 
(OL). Another colony at Bevala was active with 
breeding birds between February and March. This 
species apparently prefers to place nests in Acacia 
rovumae, which are generally the tallest trees in 
the forest and have the flattest canopy. 



24 



FIELDIANA: ZOOLOGY 



Local Names — Vorokotsy (Manombo), vorom- 
potsy (Manafiafy). 



Butorides striatus rutenbergi 
Green-backed (Striated) Heron 

The Green-backed Heron was relatively un- 
common. This species was recorded at Manafiafy, 
Itapera, near Ranopiso, Lac Anony, and Petri ky 
and along the Mandrare and Mananara rivers. It 
was relatively uncommon in the area, and most 
observations are from the edge of freshwater lakes 
and rivers and brackish lagoons. 

Breeding — A female collected at Manafiafy on 
1 1 October had a shelled egg in the oviduct. 

Weight— Female (1), 205 g. 

Soft Part Colors — Bill: maxilla black, and 
mandible black along tomia and yellow along 
base; legs: dull yellow; claws: gray; iris: yellow; 
orbital ring: yellow. 

Local Name — Keho (Manafiafy). 



Egretta ardesiaca 
Black Egret 

The Black Egret was distinctly less common in 
the area than the Dimorphic Heron. Flocks of up 
to 30 Black Egrets have been observed in rice 
paddies between Tolagnaro and Ranopiso. A sin- 
gle individual was observed along the Mananara 
River in mid-December. There are a few obser- 
vations of this bird near the RP de Berenty (MP). 



Egretta dimorpha 
Dimorphic Heron 

The Dimorphic Heron was observed in coastal 
lagoons and freshwater systems near Manafiafy, 
Itapera, Mandena, and Petri ky. To the east of the 
Anosyenne Mountains, our only records away 
from the immediate area of the coast are two in- 
dividuals in a flooded rice field near Enakara at 
about 140 m and two white-morph birds along the 
Mananara River. White-morph birds were ob- 
served at the RP de Berenty and Bevala, whereas 
dark-morph birds were occasionally seen at the 
RP de Berenty but more commonly were found 
in coastal environments. Of the 19 individuals re- 
corded during the 1989-1990 study seasons, 13 
were pure white-morph birds, two were white- 
morph birds with a few dark flight feathers, and 



four were dark-morph birds with white wing 
patches. 

Breeding — A mixed colony of this species, 
Nycticorax nycticorax, and Ardea purpurea was 
found in July at the RP de Berenty. E. dimorpha 
commenced breeding by mid-August and were on 
nests with eggs or young by 7 September. 

Local Names — Lombokoma, perhaps generic 
for large egrets (Manafiafy, Manombo); mandom 
bokomana, perhaps generic for large white ar- 
deids (Marosohy). 



Egretta alba melanorhynchos 
Great Egret 

The Great Egret was relatively common along 
lowland lakes and streams and in agricultural 
fields, particularly when flooded, throughout the 
region. It was noticeably more common east of 
the Anosyenne Mountains. Generally no more 
than three or four birds were seen together. An 
exception was several dozen birds on Lac Erombo 
on 9 July (OL & LW). 

Breeding — This species also has been found 
nesting in the RP de Berenty and has been ob- 
served along the Mananara River east of Hazo- 
fotsy. 



Ardea purpurea [madagascariensis] 
Purple Heron 

The Purple Heron was regularly noted foraging 
in areas with natural vegetation, including lake 
margins, coastal lagoons, and swamps, and irreg- 
ularly noted along the seacoast. It was markedly 
more common in marsh areas than near open wa- 
ter. Generally single birds were observed, al- 
though at dusk on 7 October, near Manafiafy, a 
group of seven birds was flushed out of a marsh. 
It was generally observed in lowland coastal ar- 
eas, but there are inland records from areas such 
as rice fields near Eminiminy and along the Man- 
drare and Mananara rivers. 

Breeding — On 24 October a pair of Purple 
Herons was observed landing in a marsh near 
Manafiafy; one bird had a stick in its bill. At the 
RP de Berenty it breeds in mixed colonies with 
Nycticorax, E. dimorpha, and Bubulcus. At this 
site A. purpurea has been seen on nests with eggs 
and juveniles in mid-October. Solitary nests have 
also been found at Bealoka. A single bird was 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



25 



flushed from a nest along the Mananara River on 
9 December. 

Local Names — Ragado (Manafiafy), rangado 
(Manombo). 



cember. We follow Kahl (1979) in our subspecific 
designation of the form occurring on Madagascar. 
Local Names — Takatry (Manombo, Maroso- 
hy), takatsy (Berenty). 



Ardea cinerea firasa 
Gray Heron 

The Gray Heron was recorded in a wide variety 
of habitats, including seashore edge, flooded ag- 
ricultural fields, lake and river margins, swamps, 
and coastal lagoons. All of our records are below 
100 m. This species was often observed with E. 
alba along the Mandrare River during the dry sea- 
son, when the river was low. At bmnh there is a 
Gray Heron specimen collected in the Tolagnaro 
area before 1891. 

Breeding — A solitary nest of this species was 
found in the RP de Berenty in an Acacia on 21 
November. 



Ardea humbloti 
Humblot's Heron 

The Humblot's Heron was rare in the region. 
We have observed this species at Lac Anony in 
January (juvenile), late July, and late December. 
The easternmmost record is a single bird on 8 
October at the mouth of the Efaho River (upper 
Lac Andriambe), just west of Tolagnaro. It has 
been reported a few times from along the Man- 
drare River near the RP de Berenty, for example 
in early July (OL & LW) and late September 
(DW). 



Scopidae 

Scopus umbretta umbretta 
Hammerkop 

The Hammerkop was relatively rare in agricul- 
tural areas below 350 m. It was occasionally ob- 
served in open areas such as along the coastal 
plain (e.g., Petriky). A vocalizing pair was ob- 
served circling over a eucalyptus grove near the 
Col de Ranopiso, opposite RNI d'Andohahela 
(parcel 3), in December. This species is also 
known from the RP de Berenty area (FO). One 
individual was observed several times along the 
Mananara River, east of Hazofotsy, in mid-De- 



Ciconiidae 

Mycteria ibis 

Yellow-billed Stork 

The only record we are aware of for the Yel- 
low-billed Stork in southeastern Madagascar is a 
flock of 15 just west of Tolagnaro (SJ). The flock 
was observed in the early morning roosting in 
some trees next to an extensive area of rice pad- 
dies. This species is known to have irregular wan- 
derings around the island (Langrand, 1990). 



Threskiornithidae 

Lophotibis cristata cristata 
Madagascar Crested Ibis 

The Madagascar Crested Ibis was relatively 
common in the majority of the forests visited from 
the Marovony Forest south through the Anosyen- 
ne Mountains to the Manantantely Forest. The 
highest elevation in the region for this species was 
recorded at the edge of the Forest of Varavara, 
not far from Ankepotsy, at approximately 1450 m. 
Despite regular human persecution, the Madagas- 
car Crested Ibis was still relatively common in the 
remaining tracts of littoral forest, particularly at 
Manafiafy, Itapera, and Mandena, and was less 
common in humid forests. We generally saw sin- 
gle birds or pairs walking on the ground, often on 
trails, and exceptionally groups of up to seven in- 
dividuals. This species will cross open areas; for 
example, in late December an adult was observed 
flying in the early morning between the Bezavona 
Forest and the littoral forest at Mandena, a dis- 
tance of about 1 km. The specimen label of a bird 
collected near Tolagnaro in 1948 notes, "common 
in dense secondgrowth forest" (fmnh). This spe- 
cies was absent from Petriky and spiny forest hab- 
itat, although it has been reported from the RP de 
Berenty (Langrand, 1990). It appears to be rare at 
the RP de Berenty; MP never observed this spe- 
cies there or at Bealoka during the course of over 
2.5 years of intensive field work in the mid-1980s. 

Local human pressure has apparently been 
present for at least 400 years (Flacourt, 1658). 



26 



FIELDIANA: ZOOLOGY 



People living near Manafiafy and Mandena have 
a variety of techniques to catch this species. One 
type of trap is made from a series of approxi- 
mately 1-m-long sticks that are inserted about 10 
cm into the ground to form a spiral funnel trap, 
the roof of which is closed off with lashed-to- 
gether Ravenala leaves, or the sticks on opposite 
sides of the structure are angled such that they 
can be attached to one another. At the closed end 
of the funnel is a chamber. Apparently, this spe- 
cies is unable to find its way back out of the trap 
from the chamber. These traps are often placed in 
areas frequented by this bird, such as narrow trails 
in the forest. Ground snares were also used in 
such places to capture this species. We found no 
evidence that any of these traps were baited. 
Snares were also used to capture adults sitting on 
nests by slowly approaching the nest and placing 
a noose attached to the end of a long stick around 
the bird's neck. We were told that nesting adults 
usually do not fly when approached. On 20 De- 
cember, under a nest in the strand forest of Man- 
afiafy, we found the plucked feathers of an adult 
Madagascar Crested Ibis. Also, at this same lo- 
cality a local resident brought us a live ibis that 
he had captured. Over the course of 4 months in 
late 1989 evidence was found near Manafiafy of 
at least five adult Lophotibis being captured and 
eaten. Nestlings and eggs are also removed from 
nests and consumed. The inaccessibility to hu- 
mans of portions of the remaining forest and the 
difficulty of finding nests are presumably the main 
factors that allow the continued existence of this 
species in the area. 

All specimens examined from the area sur- 
rounding Tolagnaro are referable to the nominate 
subspecies, including birds from Manafiafy, Man- 
dena, and "Taolanaro, Tanosy" (fmnh). Birds oc- 
curring in the RP de Berenty area may well be L 
c. urschi, the subspecies confined to dry western 
and southern forests (Langrand, 1990). 

Breeding — Near Manafiafy on 24 October an 
adult was found incubating three eggs in a nest in 
a Pandanus about 6 m off the ground, 100 m from 
the seacoast, in closed-canopy littoral forest. Nest- 
lings were noted on 10 January near Tol- 
agnaro (mnhn) and on 28 December at Mandena 
(fmnh). 

On 25 October in the strand forest of Manafia- 
fy, about 20 m from the seacoast, two birds were 
observed displaying to one another. The birds 
were flushed from the ground, and flew up and 
perched about 1 m apart on a nearly horizontal 
branch about 5 m off the ground. Periodically dur- 



ing the next several minutes they would bow to- 
wards one another, toss back their heads, return 
the bills to a horizontal position, and then touch 
bills. This display is similar to that described by 
Appert (1966). After about 10 such displays one 
bird flew off. The remaining bird moved around 
on the limb and uttered a faint, short wailing call. 
This bird, perhaps disturbed by our presence, then 
flew farther into the forest. It resumed calling 
from a new location, but this time it gave a series 
(up to 8 or 10 notes) of short, harsh notes on one 
pitch. 

In September and October we regularly heard 
this species calling during the night at various 
times between dusk and dawn. On several occa- 
sions it was observed perched in relatively tall 
dead trees, often at the forest edge or along a 
stream, where it would call almost continuously 
for up to 30 minutes. 

Diet — One individual was observed foraging in 
a small stream bed in a savanna area about 0.5 
km from the edge of the Itapera Forest. It was 
feeding on water invertebrates. In the RNI 
d'Andohahela, at 400 m, one was seen foraging 
by probing with its bill 5-10 cm into the bed of 
a forest stream. A nestling from Mandena had 
some beetle remains in its stomach, and an adult 
from near Tolagnaro had "wire worms, centipedes, 
millipedes + misc. orthop." in its stomach 
(fmnh). 

Soft Part Colors — Bill: lime green at base 
merging to olive at tip, gray with black tip (im- 
mature); legs: pinkish red, pinkish yellow (im- 
mature); claws: dull black; iris: orangish red, 
brown (immature); orbital ring: pinkish red, dull 
yellowish pink (immature). 

Local Names — Akoala (Berenty), akohola- 
hin'ala (Manafiafy, Marosohy, Marovony), ako- 
hon'ala (Manombo). 



Phoenicopteridae 

Phoenicopterus ruber [roseus] 
Greater Flamingo 

The Greater Flamingo has been recorded at a 
number of sites, and their numbers show consid- 
erable variation. There is no evidence of this spe- 
cies breeding in southeastern Madagascar. On 30 
September 11 adults and 14 immatures were ob- 
served on the lagoon near Petriky. Langrand 
(1990) noted a flock of about 3,000 on Lac Er- 
ombo in September. Records from Lac Anony in- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



27 



elude six on 7 August (OL), 180 individuals, all 
adults except four individuals, on 24 December 
(OL, LW, & SOC), and about 100, all of which 
were adults save one, on 24 December (the fol- 
lowing year). During periods of high water levels 
this species was rare or absent from Lac Anony. 
On several occasions it was observed flying over 
the RP de Berenty. 

Local Name — Sama (Manafiafy). 



Phoenicopterus minor 
Lesser Flamingo 

The Lesser Flamingo was less common than the 
Greater Flamingo in the area. On 9 July about 300 
individuals were observed on Lac Erombo (Lan- 
grand, 1990; OL & LW). These birds were in 
small groups of 5-10 individuals, mixed with 
flocks of P. ruber. During this visit flamingos 
were found for sale as food in a local market. The 
birds were captured by small nylon nooses placed 
in the shallows of the lake bottom. This same 
technique is also used in the Lac Ihotry region 
(Griveaud, 1960). Records from Lac Anony in- 
clude seven on 23 March and 59 (all adult) on 7 
August (OL & LW). 



Anseriformes 

Anatidae 

Dendrocygna bicolor 
Fulvous Whistling Duck 

The Fulvous Whistling Duck was seldom en- 
countered. A few individuals were noted along the 
Andohafasy River, near Manafiafy, and it was rare 
on the Mandrare River. 



Dendrocygna viduata 

White-faced Whistling Duck 

This species was one of the most commonly 
observed ducks in the region. The vast majority 
of records are restricted to the coastal plain. 
Flocks of up to 200 individuals were noted along 
rivers, coastal lagoons, flooded fields, and inland 
lakes. Occasionally it was found in mixed flocks 
with Anas erythrorhyncha of up to 40-50 ducks 
in total. It was occasionally found in upland areas 



up to 350 m. Observations of this species along 
the Mandrare River are occasional and sporadic. 

Breeding — In mid-December an adult and six 
downy young were observed along the Andoha- 
fasy River near Manafiafy. 

Local Name — Tsiriry (Berenty, Manafiafy, 
Manombo, Marosohy). 



Sarkidiornis melanotos 
Knob-billed Duck 

The Knob-billed Duck was not common in lit- 
toral and humid forest areas. On 14 December, 
three birds were noted at the edge of the Maro- 
sohy Forest at approximately 250 m. It also was 
noted at Manafiafy. In areas west of the Anosyen- 
ne Mountains it appears to be more regular, for 
example along the Mandrare and Mananara rivers. 
On 26 June 1984, 50 birds were observed on the 
Mandrare River between Berenty and Bealoka. 
The distinctive vocalization of this species could 
be heard in the late afternoon and at dusk over 
the Malaza and Bealoka forests. 

Local Name — Ongongo (Berenty). 



Anas melleri 
Meller's Duck 

Meller's Duck was frequently observed, but 
only in small numbers, in aquatic areas along the 
coastal plain, particularly freshwater rivers, shal- 
low lakes, and flooded grasslands up to 50 m. 
Groups of more than two individuals were seldom 
noted. 



Anas erythrorhyncha 
Red-billed Teal 

The Red-billed Teal was relatively common, 
particularly in freshwater habitats below 150 m. 
Flocks of up to 60 individuals were observed 
along rivers, lakes, and marshes and rarely in 
flooded agricultural fields. It was common in 
coastal lakes and lagoons near Manafiafy, Itapera, 
Mandena, and Petriky. Along the Mandrare River 
this species was sometimes observed with Den- 
drocygna viduata and Sarkidiornis, but always in 
small numbers. Up to 55 individuals were ob- 
served feeding and roosting along the Mananara 
River east of Hazofotsy in mid-December. On oc- 
casion large concentrations of this species are ob- 



28 



FIELDIANA: ZOOLOGY 



served, e.g., over 1,000 individuals on 9 July on 
Lac Erombo (OL & LW). 

Local Names — Boky, generic for duck (Maro- 
sohy); sadakely (Manombo). 



Anas hottentota 
Hottentot Teal 

The Hottentot Teal was uncommon. Our only 
record is two birds on 26 September at Petriky. 



Falconiformes 

Pandionidae 

Pandion haliaetus 
Osprey 

On 14 February 1988, one Osprey was ob- 
served in Tolagnaro (Langrand & Sinclair, 1994). 
This is the only known record for southeastern 
Madagascar. 



Table 1 . Observations of Milvus migrans on morn- 
ing transects in the Malaza Forest from May 1984 to 
April 1985 based on 7.3 km of trail surveyed per month. 



Month 



1984 
May 
June 
July 
August 
September 
October 
November 
December 

1985 
January 
February 
March 
April 



Total M. migrans 
observed 



20 

56 

196 

75 

49 

4 





1 

1 

16 



in flight and makes five or six flickering wing- 
beats while the wings are pointing up and down. 



Machaeramphus alcinus [anderssoni] 
Bat Hawk 



Accipitridae 

Aviceda madagascariensis 
Madagascar Cuckoo-Falcon 

In the humid forests of southeastern Madagas- 
car this species was rare. Records are from the 
Marosohy Forest (375-900 m) and the Bezavona 
Forest (75 m). The only other record from humid 
forest is three individuals displaying together over 
a ridgetop in primary forest at 810 m in parcel 1 
of the RNI d'Andohahela. This latter record was 
the only observation of this species in humid for- 
ests during the nearly 2-month expedition to the 
reserve, despite a total of 23.5 hours of canopy 
watches. In comparison, it was relatively common 
in the dry areas, particularly in gallery forest. This 
species was observed displaying on four occa- 
sions in mid-December in spiny forest near the 
Mananara River in parcel 2 of the RNI 
d'Andohahela. Other localities it has been record- 
ed include Lanirano, Andonabe, Ranomainty, Be- 
aloka, RP de Berenty, and Lac Erombo. 

The display flight of this species, noted on one 
occasion, is like that described in Safford and 
Duckworth's report (1990); a bird turns sideways 



The only records of this species are individuals 
noted on a few occasions above the Malaza Forest 
at dusk. 



Milvus migrans parasitus 
Black Kite 

The Black Kite was a common raptor in the 
open country below 100 m. We regularly ob- 
served it over river plains, open fields, and pas- 
turelands and at the edges of small villages. It was 
exceptional to find this bird within humid forest. 
This species was more common on the dry west- 
ern side of the Anosyenne Mountains than on the 
eastern side. Concentrations of Black Kites are 
not uncommon in the spiny forest region, partic- 
ularly in trees along river margins. A specimen 
(amnh) was taken by Bluntschli on 26 October 
near Amboasary-Sud. 

At certain times of the year large concentrations 
of this species were noted in the Malaza Forest 
and along the Mandrare River, and this species 
was most common during the winter months be- 
tween June and September (Table 1). Up to 30 
individuals have been found perching in a single 
tree along the Mandrare River near the RP de Ber- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



29 



enty, and hundreds may be seen at the same time 
cooling off in the river. 

Diet — At the RP de Berenty this species pro- 
vokes antipredator behavior in the Ring-tailed 
Lemur {Lemur catta) and Verreaux's Sifaka 
{Propithecus verreauxi) (Sauther, 1989; Good- 
man et al., 1994b), although there is no direct 
evidence that it attacks either species. Further, 
these two lemurs respond much less frequently 
to the passing of Milvus overhead than to Poly- 
boroides and Buteo (A. Jolly, pers. coram.). It 
has been observed at this locality taking young 
Bubulcus from nesting colonies, particularly 
those that fall from nests, and young chickens in 
nearby villages. Pellets contained beetle parts 
and larval stages of Lepidoptera. Also, it was 
observed consuming grassland invertebrates, 
particularly grasshoppers and locusts. At Lac Er- 
ombo this species was observed feeding on large 
colonial spiders and near the RP de Berenty fol- 
lowing plows in sisal plantations (OL & LW). 
The favored perches for hunting and roosting are 
Acacia rovumae, Tamarindus, Neotina isoneura, 
Quivisianthe papinae, and Albizia polyphylla, all 
of which are canopy trees. 

Breeding — In the Malaza Forest and perhaps 
throughout much of southeastern Madagascar, this 
species may breed in the austral winter; birds have 
been observed copulating in July. Further, this 
species was all but absent from the forest between 
October and February. The few birds observed 
during this latter period, which coincides with the 
breeding season of most bird species, were se- 
verely harassed by Accipiter spp., Falco newtoni, 
Eurystomus, Vanga, and Dicrurus. 

Local Names — Papango (Manafiafy), tsimala 
(Berenty). 



Polyboroides radiatus 

Madagascar Harrier Hawk 

The Madagascar Harrier Hawk was found in a 
variety of habitats throughout the region, includ- 
ing littoral, lowland, and montane forests, and in 
spiny forest from sea level to about 1000 m. It 
was less frequently observed in heavily disturbed 
open habitats, although in dry areas it was often 
seen perched in open disturbed woodland. It 
breeds in disturbed and fragmented forests sur- 
rounded by sisal plantations as long as there are 
appropriate nesting trees (e.g., Adansonia). Poly- 
boroides was not observed over humid forest dur- 
ing 23.5 hours of canopy observation in parcel 1 



of the RNI d' Andohahela but was seen in degrad- 
ed savanna habitats just outside the reserve at Is- 
aka-Ivondro and over spiny forest in parcel 2 of 
the same reserve. 

Diet — Langrand and Meyburg (1984) noted 
that in the Tolagnaro area this species forages in 
open areas on the ground, particularly around 
stones, cow dung, termite mounds, and rotten tree 
stumps. "Before completely shifting or overturn- 
ing a lump of cow dung or a stone, the bird would 
peer beneath it presumably to take cockroaches 
and beetles by surprise" (1984, p. 11). In the RP 
de Berenty Polyboroides is known to feed on fly- 
ing fox (Pteropus rufus) (Goodman & Pidgeon, 
1991) and at Bealoka on Sakalava Weavers (Plo- 
ceus sakalava). It also will pull back the bark 
from tamarind and acacia trees, hanging on with 
one leg and bracing with the wings as it probes 
crevices in trees with the other leg. This species 
also uses termite mounds as hunting perches 
(OL). 

Breeding — At least three Polyboroides nest 
sites were active in the RP de Berenty at the 
same time, where on 5 September an adult was 
observed adding sticks to a nest used the previ- 
ous breeding season, and on 23 December of the 
same year the nest contained a bird about ready 
to fledge. Nests, usually placed in baobab (Adan- 
sonia) or acacia (Acacia rovumae), are reused in 
consecutive years. Copulations have been ob- 
served in late December. The Madagascar Har- 
rier Hawk often nests close to Ploceus sakalava 
colonies, which often are in or near villages. The 
tolerance of Polyboroides by Tandroy or Mahaf- 
aly villagers could mean that rigid taboos remain 
intact or that they have a complete indifference 
to this hawk. 

Soft Part Colors — Bill: black; legs: yellow; 
iris: yellow. 

Local Names — Fihiaka (Berenty), fileliakon- 
dro (Manombo). 



Circus maillardi 
Reunion Harrier 

The Reunion Harrier has been observed only 
on few occasions in southeastern Madagascar. 
One was recorded about 5 km west of Tolagnaro 
on 27 April 1988 (DT), and a single subadult was 
noted in a marshy depression just west of Tolag- 
naro on 27 September 1990 (DWo). 



30 



FIELDIANA: ZOOLOGY 



Accipiter henstii 
Henst's Goshawk 

Henst's Goshawk was observed and heard reg- 
ularly in parcel 1 of RNI d'Andohahela in Octo- 
ber-December in the 440-m and 810-m transect 
zones. During this period, only one individual was 
seen (810 m) during 23.5 hours of canopy watch- 
es. Most individuals were detected by their dis- 
tinctive vocalization. In almost all cases (as with 
observations elsewhere in Madagascar) this call 
was heard from single birds. However, on 22 Oc- 
tober (440 m) two birds, of unknown sex, flew 
along the same line about 500 m apart, both call- 
ing loudly. 

On 25 October in parcel 1 of the RNI 
d'Andohahela, a male Henst's Goshawk appar- 
ently was following a mixed species flock, con- 
taining at least Tylas eduardi, Cyanolanius mad- 
agascarinus, Calicalicus madagascariensis, and 
Terpsiphone mutata. The hawk was climbing 
clumsily up dense vegetation on the side of a tree, 
toward the feeding flock. Many of the birds gave 
alarm calls. On sighting the observer, the raptor 
flew off. 



Accipiter madagascariensis 
Madagascar Sparrowhawk 

The Madagascar Sparrowhawk was recorded in 
the humid forests of Marovony and Marosohy be- 
tween 50 and 440 m. It also occurs in the spiny 
forest sites such as the RP de Berenty, Bealoka, 
and parcel 2 of the RNI d'Andohahela, primarily 
in gallery forests. 

Diet — On 12 December an adult Madagascar 
Sparrowhawk was netted in the Marosohy Forest 
at 375 m with a fledgling Copsychus albospecu- 
laris in its talons. At the RP de Berenty, Hypsi- 
petes madagascariensis form an important portion 
of this species largely avian diet. 

Breeding — At the RP de Berenty one pair nest- 
ed in the same Acacia rovumae tree over the 
course of 4 years. In 1983 and 1984 this nest pro- 
duced three and four young, respectively. 

Weight— [Female] (1), 285 g. 



Accipiter francesii francesii 
Frances's Sparrowhawk 

The Frances's Sparrowhawk was one of the 
most widely distributed raptors. It was found from 



sea level to about 1600 m and in a variety of 
habitats from pristine humid, gallery, and spiny 
forest, to open pastureland with small forested is- 
lands, to Eucalyptus plantations. It was noted on 
a few occasions in the city of Tolagnaro and at 
Lanirano. In 1931 an individual was collected in 
Amboasary-Sud (smf). 

Of the six individuals netted in 1989 and 1990, 
five were males and one was a female. In most 
cases it appeared that the hawk was attracted to 
the net by a struggling captured bird. Although 
our data are limited, this bias might reflect a dif- 
ference in the hunting techniques between the 
sexes, as is known for other members of this ge- 
nus (Storer, 1966). For example, males might hunt 
lower in the forest strata or feed more on small 
passerine birds. 

During the 1995 mission to the RNI 
d'Andohahela this raptor was seen at 440 m, 810 
m, and 1500 m in humid forest habitat. It was 
only seen twice at 1500 m during a total of 23.5 
hours of canopy watches. A male at 1500 m called 
from the top of an 8-m dead tree on a ridgetop at 
about 1 1 hOO. The call was a loud repeated single 
note, "whick." This same call was heard at 440 m. 

Diet — Stomachs of specimens taken in the To- 
lagnaro area contained spiders (Acari, Araneae), 
beetles (Carabidae, Scarabaeidae), grasshoppers 
(Acrididae), crickets (Gryllidae), cicadas (Cicad- 
idae), Mantodea, Odonata, ants (Formicidae), and 
reptile and bird remains (Goodman & Parrillo, in 
press). An adult male Frances's Sparrowhawk was 
captured at the Marovony Forest after it attacked 
a Zosterops tangled in a mist net. At the RP de 
Berenty A. francesii feeds on skinks and geckos, 
primarily Phelsuma, Lygodactylus, Mabuya, and 
probably Tracheloptychus, and occasionally on 
birds such as Acridotheres tristis. The capture 
technique for saurians involves flying straight to- 
ward the prey and knocking or grabbing them 
from tree trunks, and standing motionless on riv- 
erbanks and flying down to lower ground to attack 
prey. 

Breeding — A female netted on 10 October at 
Manafiafy had ovarian follicles 14 X 7 mm and 
a thickened oviduct. Males collected in September 
and in early to late October at various localities 
had testes up to 12 X 5 mm. In October an adult 
rigorously defended a presumed nesting site near 
Manafiafy and would dive-bomb passing humans, 
and on one occasion a bird actually made contact. 
A parallel type of territory defense was observed 
in the spiny forest east of Hazofotsy. At the RP 
de Berenty this species breeds in Acacia, Celtis, 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



31 



BIOLOGY LIL.^Y 
101 BURRIi I HA< i 



hhD0 6l998 



and Tamarindus trees, where they place their cup- 
shaped nests at the top of the bole where the pri- 
mary branches bifurcate; Accipiter madagascar- 
iensis generally places its nests higher up and in 
more open trees. Nestlings of A. francesii have 
been observed during late December and January. 
On 6 August a pair was noted in the gallery forest 
along the Mandrare River performing display 
flights. 

Weight— Female (1), 162 g; male (5), 111.0 ± 
4.8 (104-116) g. 

Soft Part Colors — Bill: maxilla black with a 
bluish gray tomia, and mandible black with bluish 
gray base; cere: dull greenish yellow; legs: dull 
yellow; iris: deep yellow; orbital ring: orange yel- 
low. 

Local Names — Fandraokibo (Berenty), tsipara 
(Manafiafy), tsipara korovana (Marosohy). 



Buteo brachypterus 
Madagascar Buzzard 

The Madagascar Buzzard was observed at vir- 
tually every site visited and in a variety of habitats 
from near sea level to 1950 m, including primary, 
relatively intact and degraded humid and spiny 
forest, gallery forest, and open areas such as pas- 
tureland or agricultural fields near the forest edge. 
Single individuals or pairs were generally noted. 

This species was recorded from 440 m to 1950 
m in parcel 1 of the RNI d'Andohahela, where it 
was the most common raptor. During canopy 
watches it was seen on average every 2.2 hours 
at 810 m, every 1.8 hours at 1200 m, every 1.0 
hours at 1500 m, and not at all during 5 hours at 
1900 m. No suitable canopy watchpoint was lo- 
cated at 440 m. 

Diet — In the Ankapoky Forest an adult Mada- 
gascar Buzzard was observed removing a Strep- 
topelia picturata from a mist net and consuming 
the head. At Petriky, a Madagascar Buzzard was 
noted feeding on a mouse-size lizard (probably a 
Zonosaurus). In an open agricultural area below 
the Marosohy Forest we observed an adult buz- 
zard flying off with a relatively large snake in its 
talons. At the edge of the Analalava Forest we 
noted an adult buzzard dismantling a chameleon 
between 250 and 400 mm long. The bird removed 
the head and swallowed it before alighting with 
the prey on a nest containing two young. In the 
RP de Berenty, an adult buzzard brought a young 
egret back to the nest. Both Propithecus verreauxi 



and Lemur catta give predator alarm calls when 
this raptor is heard or seen nearby. 

Breeding — On 5 November a pair of Mada- 
gascar Buzzards was found attending a nest with 
two downy young at the edge of the Analalava 
Forest. In late October of the following year the 
same nest was occupied by a breeding pair. On 
15 November in the spiny forest near Hazofotsy, 
an adult was apparently brooding the contents of 
a nest. At the RP de Berenty, this species prefers 
to nest in the canopy of tall emergent trees, such 
as Acacia rovumae, Tamarindus, and Neotina iso- 
neura, and they reuse the structure from year to 
year. At this site fledglings have been observed in 
January. A pair of Newtonia brunneicauda nested 
in the bottom of an active buzzard nest. In the 
spiny forest nests are often placed in large Al- 
luaudia trees. 

In the humid forests of parcel 1 of the RNI 
d'Andohahela at least four pairs occupied terri- 
tories along the Andranohela River between 
Trafonaomby and the forest edge near Eminiminy. 
A single individual was seen at 1950 m on 30 
November carrying a small branch. Displays 
(birds soaring high followed by steep dives and 
wingfluttering) were noted in all altitudinal zones 
within the humid and spiny forests of the reserve 
between 18 October and 12 December. 

Soft Part Colors — Downy nestling. Bill: 
black; cere: dull grayish blue, legs: off-white with 
slight bluish cast; claws: black; iris: dark tan. 

Local Names — Hindry (Andohahela), hondria 
and bevorotse (Berenty). 



Falconidae 

Falco newtoni newtoni 
Madagascar Kestrel 

The Madagascar Kestrel was one of the most 
common raptors. This species was generally re- 
corded in open areas such as agricultural fields, 
pastureland, and gardens or at the forest edge 
from near sea level to 900 m. It was seldom noted 
in areas of intact humid forest but apparently pen- 
etrates such habitat along open river channels. For 
example, our only record of this species in the 
Marosohy Forest was near a river cascade at 375 
m. F. newtoni was common in spiny forest and 
adjacent degraded savanna in parcel 2 of the RNI 
d'Andohahela. 

Diet — The stomach of a bird collected "30 km 
NNW Fort-Dauphin" contained insect remains 



32 



FIELDIANA: ZOOLOGY 



(mnhn). In the RP de Berenty this species feeds 
mostly on grasshoppers, cockroaches, locusts, and 
skinks and occasionally on small birds. 

Breeding — In the RP de Berenty F. newtoni 
often places its nest in holes in or at the top of 
broken-off trunks of Acacia rovumae and in open 
buildings and on ledges. In Tolagnaro a pair nest- 
ed on the post office tower. Adults are very vocal 
during breeding, calling to one another when re- 
turning to the nest with prey. Fledglings have 
been noted between late November and early Jan- 
uary. 

Weight— Male (1), 85 g. 

Soft Part Colors — Bill: gray at base merging 
to black tip; cere: yellow; legs: yellow; claws: 
black; orbital ring: yellow; iris: brown. 

Local Names — Rehitriky (Berenty), hitsikitsika 
(Manafiafy), hitsikitsiky (Manombo, Marosohy). 



Falco zoniventris 
Banded Kestrel 

All of our records of this species in southeast- 
ern Madagascar are confined to spiny forest, al- 
though in other areas of the island it also is found 
in humid forest (Langrand, 1990). In the Anka- 
poky Forest, the Banded Kestrel was relatively 
common. Individuals would often be noted 
perched in tall baobabs or other trees with good 
vantage points, from which they would sally out 
and capture prey in the air. This species was reg- 
ularly observed in the RNI d'Andohahela (parcel 
2), where it nests. This species also occurs in the 
RP de Berenty, Bealoka, and in nearby sisal plan- 
tations. Occasionally it is seen in open landscapes, 
perched on scattered sisal plants kilometers from 
the nearest forest. 

Diet — Two or three pellets collected below a 
perch in the Ankapoky Forest contained insect re- 
mains and the bones of at least two Ploceus sak- 
aiava. In the Malaza and Bealoka forests this spe- 
cies sometimes hunts from sisal inflorescences at 
the ecotone between forest and plantation. 

Breeding — In parcel 2 of the RNI d' Andohahela, 
an adult Banded Kestrel called in an Alluaudia 
tree for 20 minutes while holding an Oplurus liz- 
ard in one foot. The call consisted of a quiet 
"kwik . . . [repeated in a series and then ending 
with] . . . tsit-tsit-tsit," the latter three notes rising 
slightly. Another individual perched about 100 m 
away was silent. 



Falco eleonorae 
Eleonora's Falcon 

This Palearctic migrant was not common. An 
immature individual was observed on 8 December 
over parcel 2 of the RNI d'Andohahela, and an 
adult was observed on 23 December at Petriky. 
On 2 March three individuals were observed in 
parcel 3 of the RNI d'Andohahela with F. con- 
color (OL). The earliest known date of arrival in 
the region is 19 November in the Bealoka Forest. 



Falco concolor 
Sooty Falcon 

Both the Sooty Falcon and Eleonora's Falcon 
are boreal winter migrants to Madagascar. Unlike 
Eleanora's Falcon, Sooty Falcons can at times be 
relatively common in inland open areas. For ex- 
ample, on 19 November at Bealoka 12 individuals 
were observed along one path. The earliest date 
of arrival in the area is 19 October (SOC) and the 
latest date of departure is 7 May. 

Diet — The boreal autumn arrival of this species 
in the dry areas of southeastern Madagascar gen- 
erally coincided with cicada emergence. This fal- 
con perches on exposed dead trees or sisal inflo- 
rescences close to the forest edge overlooking the 
floodplain and robs sphecid wasps (Sphecius 
grandidieri) of their cicada prey by swooping at 
them. In April or May, before the falcons depart 
to northern breeding grounds, they will feed on 
Foudia madagascahensis during a period when 
invertebrate availability is declining. On 1 March 
Falco concolor was observed along the Mananara 
River, near Hazofotsy, capturing dragonflies on 
the wing (OL). 



Falco peregrinus [radama] 
Peregrine Falcon 

The Peregrine Falcon was rare. On 9 December 
one was observed in the Marosohy Forest, at the 
edge of the RNI d'Andohahela (parcel 1), at about 
375 m. From an open vantage point along the Tsi- 
tongatona River a Peregrine Falcon, perhaps the 
same individual, was noted flying above the can- 
opy and alighting on a large and slightly white- 
washed rocky outcrop high along the steep river 
valley. It is possible that the outcrop was an aerie. 
On 7 July a Peregrine Falcon was observed in the 
village of Ebelo, 60 km north of the RP de Ber- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



33 



enty (OL). Nearby is the cliff of Vohitsiombe, 
where this species might breed. Between October 
and December it was observed in the sisal plan- 
tations bordering the RP de Berenty. 

Diet — In November and December 1 986 a Per- 
egrine Falcon regularly attacked chickens in the 
village of Hazofotsy. Rand (1936) reported sev- 
eral incidences of this species feeding on domes- 
tic fowl, a prey type that is rarely taken by Per- 
egrine Falcons on the African continent (Brown 
et al., 1982). Adjacent to the Bealoka Forest this 
species has been observed on two occasions feed- 
ing on Streptopelia picturata. 



Galliformes 

Phasianidae 

Margaroperdix madagascarensis 
Madagascar Partridge 

The Madagascar Partridge was recorded at a 
variety of sites from near sea level to about 1700 
m. It was a common species, although because of 
its skulking habits in relatively dense grassland it 
was often difficult to see. When disturbed, it flew 
reluctantly and only for short distances. It was 
generally found in open grassland or heath habitat 
throughout this area, even on steep slopes. At 
least four individuals were present in degraded ar- 
eas below Trafonaomby adjacent to a marsh be- 
tween 1550 and 1700 m. 

Diet — A female caught in a mammal trap bait- 
ed with peanut butter and rice at Mandena had 
small seeds in its stomach. 

Breeding — The Mandena specimen's ovary 
was slightly enlarged (20 X 8 mm), and the larg- 
est ovarian follicle was 3 mm. 

Weight— Female (1), 250 g. 

Soft Part Colors — Bill: maxilla black, tomia 
basally bluish gray, and mandible bluish gray with 
black tip; legs: light gray with darker scute edges; 
iris: dark reddish brown. 

Local Name — Traotrao (Manafiafy, Manom- 
bo, Marosohy). 



Coturnix coturnix [africana] 
Common Quail 

The Common Quail has been reported in the 
RP de Berenty (Langrand, 1990). No other re- 



cords of it are known from southeastern Mada- 
gascar (Langrand & Appert, 1995). 

Numididae 

Numida meleagris mitrata 
Helmeted Guineafowl 

The Helmeted Guineafowl was a relatively 
common bird of open habitat, particularly damp 
marshes, grasslands, pasture, river edge, and the 
edge of degraded humid and spiny forest from 
near sea level to 325 m. It also has been regularly 
observed in sisal plantations. Occasionally large 
groups are seen, e.g., up to 40 individuals at the 
edge of gallery forest north of the RP de Berenty 
(OL). During the dry season considerable num- 
bers of this species are noted foraging in sweet 
potato fields along the Mandrare River bed. It of- 
ten visits water sources at dawn and dusk. Spec- 
imens taken at Eminiminy on 12 October (amnh) 
and 12 November at "Taolanaro, Tanosy" (fmnh) 
are referable to the form mitrata. 

This species is regularly hunted by local inhab- 
itants of the area, who use a variety of traps and 
snares to capture it. The Madagascar population 
of this species cannot be distinguished from east 
African populations, and local birds presumably 
were introduced to the island. 

Local Name — Akanga (Berenty, Manafiafy, 
Manombo, Marosohy). 



Gruiformes 

Mesitornithidae 

Mesitornis unicolor 
Brown Mesite 

The Brown Mesite was found in humid forests 
at Marovony, Analalava, Bezavona, Marosohy, 
and Manantantely from 50 to 810 m. It apparently 
prefers closed-canopy forest with a relatively 
open understory and was more common below 
100 m. This species was not detected, however, 
during the 1995 mission to the RNI d'Andohahela, 
despite 7 weeks of intensive field work in the hu- 
mid forest of parcel 1. Appert (1985) noted this 
species at Bemangidy, which is the same site 
where Hoogstraal found this species and collected 
two nests (fmnh), one on 24 November and the 



34 



FIELDIANA: ZOOLOGY 



other on 25 December. Each nest contained a sin- 
gle egg, within 2 m of the ground, and placed in 
the fork of a sloping tree (Rand, 1951). 

In late October in the Analalava Forest and in 
late September in the Manantantely Forest this 
species was very responsive to playback record- 
ings. On one occasion, recordings were played in 
an open area about 30 m from the forest edge after 
a bird was heard calling from somewhere inside 
the forest. In response, a pair of birds approached 
to within only a few meters of the recorder and 
observers, even though they were at the edge of 
the forest. Only one member of the pair called; in 
response to further playback, this bird climbed up 
a fallen log and continued calling from this perch, 
about 1 m off the ground. 

Diet — The stomach of the bird taken in the An- 
alalava Forest contained spiders (Araneae), gas- 
tropods, cockroaches (Blattodea), beetles (Chrys- 
omelidae, Curculionidae, Elateridae, Scarabaei- 
dae), and ants (Formicidae) (Goodman & Parrillo, 
in press). Appert (1985) noted that the diet of this 
species and Canirallus kioloides might be similar 
in the Bemangidy Forest, and that these two spe- 
cies might compete for resources. 

Breeding — A female collected at Analalava 
Forest on 7 November had a shelled egg in the 
oviduct and two corpora lutea. 

Weight — Female (1), 148 g (with shelled egg 
in oviduct). 

Soft Part Colors — Bill: maxilla dark brown, 
and mandible dark yellow at base merging to 
brown; legs and claws: greenish brown; iris: 
brown. 



on 30 September (amnh, smf) and reported from 
Bemangidy (Appert, 1985). Single adults were 
seen twice in open grassland adjacent to sclero- 
phyllous forest at 1700 m near Trafonaomby. 

Throughout its range this species is subjected 
to local hunting pressure. A variety of walk-in 
type traps are used to capture it, as are slingshots. 

Diet — The stomach of a bird collected at Man- 
dena had Gastropoda, Blattodea, beetles (Curcu- 
lionidae, Elateridae, Scarabaeidae, Tenebrioni- 
dae), flies (Asilidae), Hemiptera, and Lepidoptera 
(larva) remains in its stomach (Goodman & Par- 
rillo, in press). While searching for food in the 
soil or leaf litter this species makes a distinctive 
small circular or conical scrape about 9-10 cm in 
diameter, which is diagnostic of this species' local 
presence. 

Breeding — Downy young were found on 9 
September at Mandena, on 9 October at Anka- 
poky, and on 9 April at the RP de Berenty. 

Weight— Female (3), 59, 72, 80 g; male (2), 
60, 72 g. 

Soft Part Colors — Bill: dull bluish gray, light 
bluish gray, and black (downy); legs: whitish 
gray, pinkish gray blue, and brownish gray 
(downy); iris: pale cream white, light grayish 
white, and brown (downy). 

Local Name — Kibo (Berenty, Manafiafy, Man- 
ombo, Marosohy). 



Rallidae 



Rallus madagascariensis 
Madagascar Rail 



Turnicidae 

Turnix nigricollis 

Madagascar Button Quail 

The Madagascar Button Quail was recorded at 
a variety of sites from near sea level to 1700 m. 
This species was relatively common in open 
grassland, heathland, and littoral and spiny forest 
and at the edge of humid forest. It was common 
in areas that offer cover ranging from grassland 
and secondary growth with weeds to closed-can- 
opy gallery forest with thick leaf litter. Popula- 
tions continue to exist in heavily disturbed areas, 
e.g., the littoral forest near Lac Lanirano on the 
outskirts of Tolagnaro. In open grasslands this 
species was often sympatric with the Madagascar 
Partridge. Turnix were collected near Eminiminy 



Three specimens at fmnh collected at Beman- 
gidy in late November by H. Hoogstraal are the 
only known records of this species from south- 
eastern Madagascar. According to the specimen 
labels the birds were collected "in forest." 

Local Name — Kiky (Manombo). 



Dryolimnas cuvieri cuvieri 
White-throated Rail 

This species was relatively common in marsh- 
es, lake edges, or low-lying areas within or near 
to littoral and humid forest from the Marovony 
Forest south to Marosohy, Mandena, and Beza- 
vona. Appert (1985) reported this species from the 
Bemangidy Forest. It was more common in litto- 
ral forests than in upland humid forests and could 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



35 



L 



be found in close proximity to villages. The high- 
est elevation at which we recorded this species 
was in the Marosohy Forest at 900 m. During the 
1995 mission to the RNI d'Andohahela it was 
found in humid forest at 440 m in dense vegeta- 
tion along the Andranohela River but not at higher 
elevations. This species was relatively rare along 
the river margins in the region west of the Ano- 
syenne Mountains, although it is known from ar- 
eas where there is sufficient Phragmites cover 
such as along the Mananara River and near Ran- 
opiso. 

Diet — Stomachs of collected individuals con- 
tained insect remains, including Araneae and 
Scarabaeidae (Melolonthinae), and seeds (Good- 
man & Parrillo, in press). 

Breeding — Adult females collected at Man- 
dena on 1 1 September and at Manafiafy between 
9 and 13 October had ovaries up to 15 X 7 mm, 
slightly to greatly thickened oviducts, and ovarian 
follicles up to 6 mm in diameter. Two adults and 
three juveniles were found in the Malaza Forest 
on 2 1 January during a period of high water. 

Weight— Combined (5) 232 ± 34.4 (200- 
290) g. 

Soft Part Colors — Bill: distal two-thirds 
brownish gray or black merging to pink or orange 
base; legs: olive gray, brownish gray, or dark 
brown; claws: dark brown or black; iris: reddish 
orange, reddish brown, or orange crimson. 

Local Name — Tsikoja (Manafiafy). 



Canirallus kioloides kioloides 
Madagascar Wood Rail 

The Madagascar Wood Rail was found in intact 
closed-canopy humid forests at Marovony, Ana- 
lalava, Marosalohy, Marosohy, parcel 1 of the 
RNI d'Andohahela, Bezavona, and Manantantely 
between 50 and 1950 m and in the littoral forest 
of Itapera at about 20 m. It was also known from 
Bemangidy (Appert, 1985; fmnh) and Eminiminy 
(amnh). Specimens in fmnh from Analalava and 
Bemangidy are referable to Canirallus k. kiolo- 
ides. 

In the months of October through December we 
found this species to be very responsive to tape 
playback, and several times two adults followed 
by young would approach the tape recorder. In 
one case in the Marosohy Forest, at about 475 m, 
two separate pairs were called in from opposite 
directions to a site that appeared to be a territory 
boundary. 



Diet — The stomach of a collected bird con- 
tained remains of Chilopoda, Diplopoda, Hyme- 
noptera, Isoptera, and Orthoptera (Goodman & 
Parrillo, in press). 

Breeding — A male taken on 9 November in 
the Analalava Forest had testes 9X3 mm (left) 
and 4X2 mm (right). On 10 November in the 
same forest a pair was observed with two young 
that lacked white throats. In the Marosohy For- 
est, at 800 m on 30 November, two adults were 
noted with two downy young. A pair with three 
young was observed in humid forest of parcel 1 
of the RNI d'Andohahela at 700 m on 18 Octo- 
ber. 

Weight— Male (1), 172 g. 

Soft Part Colors — Bill: bluish gray proxi- 
mally merging to dull yellow tip; nasal opercu- 
lum: brownish gray; legs and claws: black; iris: 
brown. 



Sarothrura insularis 
Madagascar Flufftail 

The Madagascar Flufftail was recorded be- 
tween 20 and 1950 m in a variety of habitats, 
including grasslands (safaka) and humid forest 
(e.g., Marosohy Forest and various places in the 
RNI d'Andohahela, parcel 1). It is also known to 
occur in the savannah areas near Eminiminy and 
Vohibaka. In high-elevation zones this species 
was noted in areas of humid vegetation under 
dense canopy, sometimes considerable distances 
from running or standing water. 

Local Name — Biriky (Andohahela, parcel 1). 



Gallinula chloropus [pyrrhorhoa] 
Common Moorhen 



The Common Moorhen was infrequently en- 
countered. Most of our records are from marshes 
at the littoral forest edge and coastal lagoons. On 
a few occasions this species was observed in 
flooded rice fields in the Tolagnaro area, including 
near Mandena and Petriky. At least four pairs 
were present along a 2-km stretch of the Manan- 
ara River in parcel 2 of the RNI d'Andohahela in 
December. They only occurred in areas with 
dense reed beds. 

Local Name — Taleva (Marosohy). 



36 



FIELDIANA: ZOOLOGY 



Porphyrula alleni 
Allen's Gallinule 

On 30 September 1990 a single adult Allen's 
Gallinule in breeding plumage was observed west 
of Tolagnaro in a marsh near the Ilot des Portugais 
(DWo). This is the only known record from the 
area. 



Porphyrio porphyrio [madagascariensis] 
Purple Swamphen 

The Purple Swamphen was relatively uncom- 
mon. The vast majority of our records are from 
large marshes adjacent to littoral forest, e.g., at 
Manafiafy and Mandena. This species also has 
been reported in the RP de Berenty (Langrand, 
1990). 

Local Name — Talevana (Manafiafy, Manom- 
bo, Marosohy). 



Fulica cristata 
Red-knobbed Coot 

The Red-knobbed Coot was noted on a few oc- 
casions. Observations include five individuals on 
9 July on Lac Erombo and one individual on 1 
August on a pond between Lac Erombo and Lac 
Anony (OL & LW); most of these records in- 
volved birds with bright red casques. 



Charadriiformes 

Jacanidae 

Actophilornis albinucha 
Madagascar Jacana 



Rostratulidae 

Rostratula benghalensis [benghalensis] 
Greater Painted Snipe 

The Greater Painted Snipe was only recorded a 
few times. On 31 October one bird was noted near 
dusk in a rice field just below Antseva (800 m), 
on 10 October several individuals were observed 
after dark in a flooded field near Manafiafy, and 
on 7 October one bird was recorded along the 
Mandrare River at the RP de Berenty. 



Glareolidae 

Glareola ocularis 

Madagascar Pratincole 

The Madagascar Pratincole, a migratory spe- 
cies that spends the austral winter in East Africa, 
was observed on numerous occasions in lowland 
areas between Bemangidy and Mandena. Our 
earliest record is from 22 September at Pointe 
Evatra. Generally small groups were observed in 
open fields or "heathland," often in areas with 
rocky outcrops. The largest concentration ob- 
served in the region was a flock of 15 birds on 
28 October resting on a rocky outcrop 40 km 
south of Manantenina. Between 14 and 16 Oc- 
tober, two pairs were observed resting on a sandy 
island in a coastal lagoon near Itapera. No nests 
were located in the region, but the birds appar- 
ently were territorial and were suspected of 
breeding. In early October Bluntschli found this 
species near Eminiminy (approximately 300 m) 
and collected several specimens (amnh, nhb, 
smf). 

Soft Part Colors — Bill: blackish brown; legs: 
black; iris: reddish brown. 



The only record we have from the area is two 
males taken near Amboasary-Sud on 22 October 
1931 by Bluntschli (amnh, smf). This is a consid- 
erable range extension; the closest known previ- 
ous locality for this species was in southwestern 
Madagascar (Langrand, 1990). We know of no 
more recent evidence of this species in southeast- 
ern Madagascar. 

Soft Part Colors — Bill: bluish gray; legs: 
bluish gray; iris: olive brown. 



Charadriidae 

Pluvialis squatarola 
Black-bellied Plover 

This migrant from Eurasia was noted on several 
occasions along the seacoast. The majority of our 
records are from the Manafiafy area, and the ear- 
liest date of arrival is 10 September. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



37 



Charadrius hiaticula [tundrae] 
Common Ringed Plover 

This Eurasian migrant was relatively common 
along the sandy beaches from Manantenina to To- 
lagnaro. Our earliest date of arrival was mid-Oc- 
tober. Flocks of up to a few hundred were ob- 
served. Tundrae probably is the form visiting the 
region. 

Local Name — Salaly kely, generic for small 
shorebird (Manafiafy). 



Charadrius tricollaris [bifrontatus] 
Three-banded Plover 

On 1 November an adult Three-banded Plover 
was observed at the edge of the Marovony Forest 
along a streambed in a grassland area. The bird 
was feeding in the shallows of the stream at the 
edge of a rock cascade. It also has been noted on 
7 July along the dry margins of the Mandrare Riv- 
er, near the RP de Berenty (OL). In general this 
species is rare in eastern Madagascar (Langrand, 
1990). 



Charadrius thoracicus 
Madagascar Plover 

This species was observed in the coastal lagoon 
near Itapera. In mid-October 1990 at least eight 
pairs were noted in the area. 

Breeding — A downy young was collected on 
19 October at the edge of Lac Mananivo near 
Itapera. 

Soft Part Colors — Downy young. Bill: base 
lime green merging to brownish black tip; legs: 
dull lime green; claws: black; iris: brown; skin 
color over most of body: slate to black. 



Charadrius leschenaultii 
Greater Sand Plover 

The Greater Sand Plover was observed on a 
few occasions, generally along the seacoast or 
slightly inland at lakes or lagoons. The earliest 
records are from early October. Flocks of up to 
15 individuals have been recorded at sites such as 
Mandena, Lac Anony, and Lac Andriambe. 



Charadrius marginatus tenellus 
White-fronted Plover 



Charadrius pecuarius pecuarius 
Kittlitz's Plover 

Kittlitz's Plover was relatively common below 
50 m elevation in grassland or sandy areas adja- 
cent to fresh or saline lakes and marshes, e.g., 
Manafiafy, Petriky, and Lac Anony. On several 
occasions we found this species in recently 
burned grassland fields, and it appears to be one 
of few ground-dwelling species that utilizes this 
habitat. Kittlitz's Plover was the most common 
small plover on the lower Mandrare River. It con- 
gregates along the shores of Lac Anony during 
seasonal periods of low water. Bluntschli collect- 
ed three specimens at Amboasary-Sud on 10 No- 
vember (amnh, smf). 

Breeding — On 16 October downy young of 
this species were found at the unvegetated edge 
of a lagoon near the Itapera Forest. At this same 
site C. thoracicus also was nesting, but this spe- 
cies appeared to prefer the slightly higher areas 
adjacent to the lagoon being used as cattle pas- 
ture. 

Soft Part Colors — Downy young. Bill: black; 
legs: yellowish brown; iris: brown. 



The White-fronted Plover was rare. This spe- 
cies has been observed at Lac Anony on numer- 
ous occasions, where it tended to occur in non- 
flooded sandy areas, often mixed in with groups 
of C. pecuarius. 

Diet — The stomach contents of four specimens 
taken at Lac Anony contained marine crustaceans 
and fine stones (PBZT). 



Scolopacidae 

Limosa lapponica lapponica 
Bar- tailed God wit 

The Bar-tailed Godwit, a Eurasian migrant to 
Madagascar, was uncommon during the boreal 
winter along coastal beaches. One individual, re- 
ferable to the nominate form, was collected in 
mid-October at Pointe Evatra (fmnh). The bird 
was killed by a local boy with a slingshot. Our 
earliest date for the region was 7 October at Man- 
afiafy. 

Local Name — Salaly be, generic for large 
shorebirds (Manafiafy). 



38 



FIELDIANA: ZOOLOGY 



Numenius phaeopus 
Whimbrel 

The Whimbrel, a Eurasian boreal winter mi- 
grant to Madagascar, was relatively common, but 
never in groups exceeding three individuals, along 
coastal beaches (e.g., Manafiafy, Itapera, and 
Pointe Evatra) and lakes (e.g., Lac Anony). We 
have no far-inland records of this species from 
southeastern Madagascar. It was observed be- 
tween late September and late October. 



Numenius arquata [orientalis] 
Eurasian Curlew 

This Eurasian migrant to Madagascar during 
the boreal winter was observed on a few occa- 
sions along the seacoast near Itapera and Mana- 
fiafy. The earliest records are from late September. 



margins in the spiny forest, e.g., one on 2 March 
along the eastern edge of parcel 3 of the RNI 
d'Andohahela and up to three individuals in mid- 
December along the Mandrare and Mananara riv- 
ers in parcel 2 of the same reserve. 



Arena ria interpres 
Ruddy Turnstone 

The Ruddy Turnstone was relatively common 
along the seacoast, particularly in rocky areas 
(e.g., Manafiafy), between early October and late 
December. This migrant from Eurasia is a boreal 
winter visitor to Madagascar. It was generally ob- 
served singly or in groups of up to three individ- 
uals, although concentrations have been noted at 
Lac Anony on 20-21 September and 24 Decem- 
ber. Presumably the nominate form is the one oc- 
curring in the southeastern region. 



Tringa nebularia 

Common Greenshank 

Small numbers of Common Greenshank were 
noted on several occasions between September 
and December. This Eurasian migrant to Mada- 
gascar during the boreal winter was generally ob- 
served on inland freshwater lakes or coastal la- 
goons, e.g., Manafiafy, Itapera, and Mandena. 
This species and the next are the two most com- 
mon species of waders in southeastern Madagas- 
car; during this period they are also found on in- 
land rivers such as the Mandrare and Mananara. 
The largest reported concentration of Common 
Greenshanks in the area was over 200 individuals, 
at Lac Anony on 24 December (OL, LW, & SOC). 
On 10 November Bluntschli collected one near 
Amboasary-Sud (amnh). 






Actitis hypoleucos 
Common Sandpiper 

The Common Sandpiper is a migrant from Eur- 
asia during the boreal winter. It was recorded in 
a variety of habitats: along the edge of the sea, 
shores of lagoons, and inland lakes, flooded 
grassy areas, and rice paddies from sea level to 
about 450 m. Records of this species in the region 
span the period from mid-October to late March. 
Generally single birds or pairs were observed. Oc- 
casionally, this species was found along river 



Gallinago macrodactyla 
Madagascar Snipe 

The Madagascar Snipe was rarely observed. 
Our records include displaying birds at Mandena 
over a small marsh in September and several in- 
dividuals at the edge of the Analalava Forest in a 
damp grassland in early November. A specimen 
was collected at "Taolanaro, Tanosy" on 14 No- 
vember (fmnh). 

Local Name — Arakarandroka (Manombo). 



Calidris alba 
Sanderling 

The Sanderling, a boreal winter migrant to 
Madagascar, was regularly observed along the 
seacoast between Manafiafy and Tolagnaro. The 
earliest record was a few at Lac Anony on 20-21 
September. The largest recorded local concentra- 
tion was a flock of over 50 birds on 25 December 
at Lac Andriambe. This species was rare along 
the Mandrare River near the RP de Berenty. 



Calidris ferruginea 
Curlew Sandpiper 

The Curlew Sandpiper is a boreal winter mi- 
grant to the island. It was recorded on numerous 
occasions, particularly along coastal beaches, la- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



39 



goons, and lakes. The vast majority of our records 
fall between late September and late December, 
although there are reports from the austral winter 
months. Concentrations of this species are occa- 
sionally noted, for example about 100 individuals 
along the shore of Lac Anony in late December. 



Stercorariidae 

Catharacta antarctica 
Subantarctic Skua 

An individual banded on Marion Island 
(46°55'S, 37°45'E) was recovered at Tolagnaro in 
March 1987 (Langrand, 1990). 



e.g., 35 on 25 December 1992 at Lac Andriambe. 
We found no evidence of this species breeding 
locally. 



Sterna caspia 
Caspian Tern 

The Caspian Tern was relatively uncommon. 
We observed this species along the seacoast near 
Manafiafy and Lac Anony in November and De- 
cember. Generally single individuals or pairs were 
noted. This species also has been observed inland 
along the Mandrare River near the RP de Berenty 
(OL & LW). 



Stercorarius longicaudus 
Long-tailed Skua 

The only record of this species in southeastern 
Madagascar is a group of four birds observed off 
the coast of Tolagnaro on 22 December 1991 
(Langrand & Sinclair, 1994). 



Sterna hirundo 
Common Tern 

On 22 August 1990, 23 Common Terns were 
observed off the coast of Tolagnaro (Langrand & 
Sinclair, 1994). This is the only record for this 
species in southeastern Madagascar. 



Laridae 

Larus dominicanus 
Kelp Gull 

The Kelp Gull was regularly observed along 
the coast near Tolagnaro, north to Manafiafy and 
west to Lac Anony. Generally there was a pre- 
ponderance of adults in the area. For example, on 
24 December 1992 a flock of 36 individuals, all 
adults, was observed at Lac Anony. This gull oc- 
casionally flies up the Mandrare River as far as 
the RP de Berenty. There is no evidence of this 
species breeding in southeastern Madagascar, al- 
though it is present in varying numbers through- 
out the year. 

Local Name — Kolokoloky (Berenty area). 



Sternidae 

Chlidonias hybridus [sclateri] 
Whiskered Tern 

The Whiskered Tern was observed several 
times along inland lagoons between late Septem- 
ber and late December. Flocks have been noted, 



Sterna dougallii 
Roseate Tern 

The Roseate Tern was observed on several oc- 
casions off the coast of Manafiafy and Tolagnaro 
and near Lac Andriambe. 



Sterna sandvicensis 
Sandwich Tern 

Two Sandwich Terns were observed on 22 Au- 
gust 1990 off the coast of Tolagnaro (Langrand & 
Sinclair, 1994). This is the only known record of 
this species on Madagascar. 



Sterna bergii 

Greater Crested Tern 

Flocks of Greater Crested Terns were observed 
numerous times along the seacoast, generally be- 
tween Manantenina and Tolagnaro and west to 
Lac Anony. In many cases the birds were a con- 
siderable distance offshore, and it was often dif- 
ficult to determine if the flocks were composed of 
this species, the Lesser Crested Tern, and/or the 



40 



FIELDIANA: ZOOLOGY 



Roseate Tern. We found no evidence of any Ster- 
na spp. breeding in southeastern Madagascar. 



Local Name — Hatrakatraka or hatrakatra 
(Bealoka, Berenty, Hazofotsy). 



Sterna bengalensis 
Lesser Crested Tern 

The Lesser Crested Tern was regularly ob- 
served off the coast, particularly in the region be- 
tween Manafiafy and Tolagnaro. Flocks of up to 
30-40 individuals were often noted foraging in 
the area of the sea where the waves started to 
break. The form occurring in the region is pre- 
sumably S. b. par. This species seemed more 
common than the Greater Crested Tern, but in nu- 
merous cases crested terns far offshore could not 
be identified to species. 



Columbiformes 

Pteroclididae 

Pterocles personatus 
Madagascar Sandgrouse 

The Madagascar Sandgrouse is an inhabitant of 
open areas in the spiny forest. We know of no 
records from east of the western slopes of the An- 
osyenne Mountains; the eastern limit of this spe- 
cies appears to be near Mahamavo and Tanambao- 
Ankatsaky and slightly below the western limit of 
parcel 1 of the RNI d'Andohahela. This species 
was regularly observed near Hazofotsy and other 
areas within and near parcel 2 of this reserve. In 
the Ankapoky Forest flocks of two to six individ- 
uals were noted flying over in the early morning. 
Delacour's (1932, p. 33) statement that "II existe 
probablement d'Analalava a Fort Dauphin" can- 
not be substantiated. 

This species appears to have a preference for 
feeding in slightly degraded or open spiny forest 
or areas dominated by Xerophyta or introduced 
grasses around rocky ground. Flocks of up to 40 
individuals often visit such sites on a daily basis 
in the late afternoon, e.g., the area near the Ha- 
zofotsy tombs. In early morning, particularly dur- 
ing the dry season, flocks of up to 100 individuals 
fly to the Mandrare and Mananara rivers to drink. 
In December this species was observed visiting 
and drinking from a tributary of the Mananara 
River throughout most of the day. 



Columbidae 

Streptopelia picturata picturata 
Madagascar Turtle Dove 

The Madagascar Turtle Dove was the most 
common columbid in the humid portion of the 
region from the Marovony Forest south along the 
coast and on the east side of the Anosyenne 
Mountains to Petriky. It occurs in intact and par- 
tially disturbed humid and littoral forests, heavily 
degraded forest, eucalyptus plantations, gardens, 
and agricultural fields from sea level to 1950 m. 
West of the Anosyenne Mountains, in transitional 
forest and spiny forest, it is the second most fre- 
quently encountered dove after Oena capensis. 
Streptopelia was common in gallery forest along 
the Mandrare River. 

Diet — The stomachs of four individuals con- 
tained seeds. These birds feed predominantly on 
forest trails, paths, or dirt roads, from which they 
pick up seeds and grains. In humid forest they are 
usually seen walking around on forest litter. In 
November at the higher elevations of parcel 1 of 
the RNI d'Andohahela this species was often ob- 
served feeding on the seeds of Sloanea. One col- 
lected bird had 15 Sloanea seeds in its crop, all 
lacking the fleshy orange aril. 

Breedincj — This species appears to have a rel- 
atively long breeding season. Active nests with 
eggs were found in the RP de Berenty on 3 July 
and 5 September (OL). In both cases the nests 
were less than 4 m off the ground and placed in 
a slender tree or in a thick bush. On 22 October 
in the Manafiafy Forest, an adult was attending a 
nest with one or two eggs. Adult males collected 
between mid-September and late December had 
testes up to 17 x 7 mm (left) and 12 x 5 mm 
(right), and adult females had ovaries up to 10 X 
6 mm, enlarged oviducts, and ovarian follicles 7 
mm in diameter. A female collected on 22 No- 
vember in parcel 1 of the RNI d'Andohahela was 
in reproductive condition and had well-developed 
"milk glands" within the crop. A fledgling was 
netted at 1500 m on 21 November in parcel 1 of 
the RNI d'Andohahela. 

Weight— Female (6), 191.2 ± 12.3 (167-200) 
g; male (3), 190, 190, 192 g; combined (12), 
190.8 ± 12.8 (165-210) g. 

Soft Part Colors — Bill: generally gray at 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



41 



1 



base, distally pale bluish gray or light greenish 
gray, and gray tip, but in two adult males in 
breeding condition crimson at base; legs: dorsally 
dull pinkish red or purplish red and ventrally light 
red, greenish gray, or bluish gray; iris: brown, 
purplish red, or red; orbital ring: purplish red, 
crimson, dull red, or dark vermilion. 

Local Names — Deho (Berenty, Manafiafy, 
Marosohy), kimokimo (Manombo). 



Oena capensis aliena 
Namaqua Dove 

The Namaqua Dove was common to abundant 
in areas of spiny forest; it was one of the more 
regularly netted species in the Ankapoky Forest. 
From the Anosyenne Mountains eastward this 
species was distinctly less common and generally 
confined to coastal areas. We noted it near Man- 
antenina, Mandena, Tolagnaro, and Petriky. There 
are few inland records from the humid forest 
zone. This species has been observed near Isaka- 
Ivondro on several occasions, mostly during the 
dry season. Bluntschli collected three Namaqua 
Doves near Eminiminy between 15 October and 
1 November (amnh, smf). The elevational range 
of this species is from near sea level to 100 m. 

Diet — This species utilizes sand or dirt roads, 
trails, and paths in open areas to search for food 
and is not found in closed canopy or dense forest 
habitat. It feeds primarily on seeds. 

Breeding — Several males netted at Ankapoky 
in mid-October had testes up to 12 X 19 mm, and 
females had large ovaries and ovarian follicles up 
to 20 mm. 

Weight— Female (3), 35, 36, 38 g; male (5), 
38.4 ± 2.5 (36-42) g; combined (10), 37.9 ± 2.51 
(36-42) g. 

Soft Part Colors — Bill: maroon at base, or- 
ange in middle, and dull yellow or light orange 
tip (males) and dull reddish brown or brownish 
black (females); legs: maroon, gray, or pinkish 
red; claws: gray or black; iris: brown. 

Local Names — Tsikaloto (Berenty). 



Treron australis australis 
Madagascar Green Pigeon 

The Madagascar Green Pigeon was a relatively 
common inhabitant of intact and degraded humid 
and littoral forests from the Marovony Forest 
south along the coastal plain and through the An- 



osyenne and Vohimena mountains to Tolagnaro, 
from sea level to about 1000 m. On a few occa- 
sions flocks of up to 20 individuals were seen fly- 
ing around gardens in Tolagnaro, presumably ex- 
ploiting ripening fruits. It was uncommon in spiny 
forest, where it may be only a seasonal visitor. 
One was taken by Bluntschli on 17 November at 
Amboasary-Sud (amnh). In 7 weeks of field work 
between October and November in parcel 1 of the 
RNI d'Andohahela, this species was only noted 
once (at 440 m), which suggests that at this season 
fruits eaten by this species were not common in 
this humid forest sector or that in this region it 
does not occupy extensive areas of closed-canopy 
forest. 

Diet — A large component of this species' diet 
appears to be Ficus fruits, and its movements may 
be related to local availability of this resource. 
One Madagascar Green Pigeon collected from a 
flock of 12 feeding in a Ficus tree at Manafiafy 
had 65 fruits in the crop that in total weighed 41 
g. At Itapera this species was observed feeding on 
Abrus fruits. In the Marovony Forest three indi- 
viduals were observed with two Alectroenas mad- 
agascariensis consuming Uapaca fruits. At Lan- 
irano it was often noted eating mulberry (Moms) 
fruits. In the RP de Berenty it was seen feeding 
on the ripe and unripe small fruits of Ficus grevei 
and Phyllanthus seyrigii. 

Breeding — A female taken on 24 October at 
Manafiafy had a slightly enlarged ovary, thick- 
ened oviduct, and ovarian follicles up to 6 mm in 
diameter. 

Weight— Combined (2), 215, 215 g. 

Soft Part Colors — Bill: vermilion at base 
with horn-color tip or crimson at base with cold 
gray tip; legs: dull orange or orangish yellow; 
claws: black; iris: cobalt blue or bluish white. 

Local Names — Bohaky (Manafiafy, Maroso- 
hy), fonomavo (Manombo). 



Alectroenas madagascariensis 
Madagascar Blue Pigeon 

The Madagascar Blue Pigeon was noted in a 
variety of habitats from intact to heavily disturbed 
humid and littoral forest from Marovony south 
through the Vohimena and Anosyenne mountains 
to just north of Tolagnaro at elevations from near 
sea level to 1950 m. The locality of a specimen 
taken in 1756 was listed as Fort-Dauphin (Stre- 
semann, 1952). One was collected at Eminiminy 
on 27 October by Bluntschli (smf). This species 



42 



FIELDIANA: ZOOLOGY 



will often sit in the top of canopy trees, particu- 
larly emergents with dead wood, to vocalize. 

Diet — The stomach of an individual taken 7 
km north of Tolagnaro on 8 May contained seeds 
of an unidentified plant locally called rithala 
(mnhn). In the littoral forest of Manafiafy this spe- 
cies was observed feeding on fruits in a Ficus 
alongside Hypsipetes and Coracopsis nigra. In 
early November Alectroenas, Treron, and Eule- 
mur fulvus were regularly observed feeding on 
fruiting Uapaca trees in the Marovony Forest. 

Breeding — The specimen collected by 
Bluntschli had small testes. In parcel 1 of the RNI 
d'Andohahela this species was noted nest building 
at 1200 m on 12 November and at 1500 m on 23 
November. The nest consists of a shallow plat- 
form of twigs placed about 3 m off the ground in 
small shrubs. Both adults participated in nest con- 
struction and flew 10-200 m away from the site 
to collect sticks. On returning to the nest site, the 
adults regularly gave a low, muted cooing call. 

Soft Part Colors — Bill: dark green; legs: 
black; iris: red. 

Local Names — Foli manga (Manafiafy), foly 
manga (Marosohy), fonomity (Manombo). 



Psittaciformes 

Psittacidae 

Coracopsis vasa vasa and [Coracopsis vasa 
drouhardi] 

Greater Vasa Parrot 



forest of parcel 2 of the RNI d'Andohahela, C. 
vasa was about equally common as C. nigra. 

A specimen collected in the Bemangidy Forest 
on 26 December is referable to C. v. vasa (fmnh). 
This is presumably the form occurring from the 
Marovony Forest south through the Anosyenne 
and Vohimena mountains. West of the Anosyen- 
nes, in the spiny forest, C. v. drouhardi is the 
expected subspecies. 

Diet — In an open area at the edge of the Ma- 
rovony Forest three Greater Vasa Parrots were ob- 
served feeding in a Ficus tree. In gallery forest 
this parrot often feeds on Tamarindus fruits. A 
flock of at least 10 birds was observed in Bealoka 
on the forest floor feeding on fallen pods. It also 
was observed in the same region consuming the 
fruits and seeds of Ficus megapoda, Celtis gom- 
phophyla, C. phillipensis, Quivisianthe papinae, 
and Neotina isoneura and the leaves of Acacia. 

Breeding — Displaying C. vasa in parcel 2 of 
the RNI d'Andohahela in December showed bare 
orange skin on their heads. It is not clear if the 
lack of feathering is caused by the mechanical or 
hormonal aspects of courtship — the male grasps 
hold of the head of the female and jerks it up and 
down in display, which might cause head feathers 
to be lost. However, apparent male birds were 
seen with this feature, suggesting that it is not 
only caused by mechanical methods. Information 
on the breeding display and copulation behavior 
of both Malagasy Coracopsis in captivity have 
been reviewed by Wilkinson (1990) and Wilkin- 
son and Birkhead (1995). 

Local Names — Kia (Marosohy); vazambe (Be- 
aloka, Berenty); boeza, generic for Coracopsis 
spp. (Manombo). 



The Greater Vasa Parrot was generally uncom- 
mon. It was observed in intact and degraded hu- 
mid forests and in agricultural areas on lateritic 
soils, e.g., in or at the edge of the Marovony, An- 
alalava, Bemangidy, and Marosohy forests. It was 
relatively common in spiny forest and in sisal 
plantations. This species' elevational range is 
from near sea level to 1950 m. Previously this 
species was thought to occur only to about 1000 
m (Langrand, 1990). There is no documented rec- 
ord of it in littoral forest. This species was less 
common than but broadly sympatric with C. ni- 
gra. Flocks of C. vasa are seldom larger than 
three or four individuals. Between October and 
December in parcel 2 of the RNI d'Andohahela, 
point count contact frequency of C. vasa was only 
20% of the frequency of C nigra. In the spiny 



Coracopsis nigra nigra and [Coracopsis ni- 
gra libs] 

Lesser Vasa Parrot 

The Lesser Vasa Parrot was relatively common 
throughout the region, including in intact and de- 
graded humid, littoral, gallery, and spiny forest, 
agricultural areas, sisal plantations, and gardens in 
villages. It was recorded from sea level to 1800 
m. Flocks of 10 individuals were not uncommon, 
and one group of 19 birds was noted at the edge 
of the Marovony Forest. This species is locally 
considered an agricultural pest, particularly with 
ripening corn and rice crops, and is consequently 
persecuted by local people. It is generally far 
more numerous and vocal than C. vasa (Table 2). 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



43 



Specimens from Manafiafy and Eminiminy are 
referable to C. n. nigra, which is presumably the 
form occurring in the Anosyenne and Vohimena 
mountains east to the coast, whereas the popula- 
tions west of the Anosyenne Mountains in spiny 
forest may be libs. 

Diet — The stomach of a specimen taken 7 km 
north of Tolagnaro on 19 February (mnhn) con- 
tained ripe fruits of a plant locally known as ro- 
hala. In the Marosohy Forest at about 900 m a 
flock of six Lesser Vasa Parrots was feeding on 
the fruits of a Macaranga. This parrot species also 
has been observed feeding on Sarcolaena multi- 
flora, almost open buds of Symphonia and Hum- 
bertia madagascariensis, the blue pericarps of 
Ravenala madagascariensis, and the seeds or 
fruits of Tamarindus, Alluaudia procera, Agava 
rigida, Celtis phillipensis, Hippocratea rubignosa, 
Acacia farnessiana, Croton, Ficus, Melia azedar- 
ach, Uapaca, and Neotina isoneura. 

Breeding — An adult male taken on 23 October 
had small testes. The specimen from 7 km north 
of Tolagnaro had a slightly enlarged ovary. 

Weight— Combined (2), 215, 220 g. 

Soft Part Colors — Bill: brownish horn; legs: 
dark brown or bluish gray; claws: gray or black; 
iris: dark brown. 

Local Names — Boloky (Manafiafy), kia (Ma- 
rosohy), vaza (Manafiafy), vazatsihotsy (Berenty). 



Agapornis carta carta and Agapornis carta 
ablactanea 

Gray-headed Lovebird 

The Gray-headed Lovebird was relatively com- 
mon in spiny forest and distinctly less common in 
agricultural and grassland areas near and in humid 
forest from sea level to 325 m. In the spiny bush 
it can often be seen in sisal plantations and 
perched on dead trees or Alluaudia procera. This 
species was distinctly uncommon in littoral forest, 
e.g., Mandena and Itapera. There is a specimen 
collected in 1756 near Fort- Dauphin (Stresemann, 
1952). It was often noted in flocks ranging in size 
from 3 to 20 individuals. The nominate subspecies 
is found from the Marovony area south through 
the Vohimena Mountains, at least to the eastern 
slopes of the Anosyenne Mountains and presum- 
ably south toward the coastal forest, and ablac- 
tanea is found from the western side of the An- 
osyenne Mountains to the Mandrare River. A 
specimen taken by Milon 30 km NNW Fort-Dau- 
phin is referable to A. c. cana (mnhn), whereas 



Table 2. Observations of Coracopsis vasa and C. 
nigra on morning transects in the Malaza Forest from 
May 1984 to April 1985, based on 7.3 km of trail sur- 
veyed per month. 





Total observed 


Month 


C. vasa 


C. 


nigra 


1984 








May 
June 


3 
7 




6 



July 
August 
September 
October 


2 

5 





8 
11 

14 
7 


November 


3 




8 


December 







7 


1985 








January 

February 

March 




2 

2 




20 
26 
12 


April 


2 




13 



specimens collected at Ankapoky are referable to 
A. c. ablactanea (fmnh). 

Diet — The crop and stomach of the Milon 
specimen contained many grass seeds. This spe- 
cies visits the heads of sisal flower stalks and will 
also descend to ground to feed on fallen seeds. 
Foraging flocks are often more active in the morn- 
ing than in the afternoon and consist dispropor- 
tionately of males (Table 3). 

Breeding — All four adults taken in mid-Octo- 
ber in the Ankapoky Forest had small reproduc- 
tive organs; one immature with a bursa also was 
collected. The Milon specimen taken on 26 May 
had testes 4.5 X 2.2 mm. On 1 March, in the RP 
de Berenty, a female was observed in gallery for- 
est entering a hole in a tree about 5 m off the 
ground (OL); nesting sites include holes in dead 
Tamarindus, Acacia, Neotina, Alluaudia, and 
Quivisianthe trees. Nesting apparently commenc- 
es in April or May. Chicks fledge and sites are 
vacated before the arrival of Eurystomus glaucu- 
rus in October, which also occupies the same 
holes. If a site is not used by Eurystomus, then 
lovebirds will occupy and visit holes during por- 
tions of the nonbreeding season. The timing of 
this parrot's breeding season may be an adaptation 
to avoid competition for nesting sites with the 
larger and more aggressive Eurystomus. 

Weight— Combined (5), 29.8 ± 1.9 (27.0- 
31.5) g. 

Soft Part Colors — Bill: flesh gray or cold 



44 



FIELDIANA: ZOOLOGY 



Table 3. Observations of Agapornis cana on morning and evening transects conducted in the RP de Berenty 
May-September (14.6 km of trail per month) and October-April (7.3 km of trail per month). 













Number 




% 






Number 








of trees 


% 


trees 


% 




of birds 


Largest 


Sex ratio 


Trees 


used as 


trees 


Tamar- 


trees 


Month 


• III! [Ill 1 


flock 


(<$:$) 


visited 1 


perches 


dead 


indus 


Acacia 


1984 


















May 


30/4 


15 


27.5 


At, Ab, K, 
Qu, K 


16 


25 


13 


38 


June 


53/16 


10 


8.3 


At, K 


7 


43 


29 


29 


July 


69/26 


9 


29.7 


Al, At, Co, 
Cr, K, Qu, S 


13 


54 


23 


15 


August 


72/— 


9 


25.7 


At, K, S 


6 


50 


17 


50 


September 


58/2 


8 


3.1 




1 


100 


— 


— 


October 


48 


9 


2.0 


At, K 


3 


33 


33 


33 


November 


30 


4 


4.1 


Ap, At. K, Qu 


8 


13 


63 


13 


December 


15 


7 


2.0 


Al, At 


2 


— 


— 


50 


1985 


















January 


37 


10 


4.2 


At, K 


4 


50 


25 


25 


February 


49 


11 


7.5 


K 


1 


— 


100 


— 


March 


26 


11 


1.1 


At 


2 


50 


— 


50 


April 


4 


4 


— 


— 


— 


— 


— 


— 


Average 


— 


8.9 


6 = 71.4% 




— 


36.5 


25.4 


28.6 



1 Ab = Acacia farnessiana, Al = Alluaudia procera, Ap = Albizia polyphylla. At = Acacia rovumae, Co = Croton, 
Cr = Crataeva excelsa, K = Tamarindus indica, N = Neotina isoneura, Qu = Quivisianthe papinae, S = Agava 
rigida. 



grayish white; legs: gray or bluish gray; claws: 
black; iris: brown. 

Local Names — Fahvaza (Berenty), kanaka 
(Berenty, Marosohy), kariaky (Manafiafy), kitreo- 
ky (Manombo), sarivazy (Manafiafy). 



Cuculiformes 

Cuculidae 

Cuculus rochii 

Madagascar Lesser Cuckoo 

The Madagascar Lesser Cuckoo migrates to 
East Africa during the austral winter (May-Au- 
gust). When this species arrives in southeastern 
Madagascar it is practically ubiquitous. We noted 
it calling for hours at a time, often at night, from 
high perches in or at the edge of humid, gallery, 
and spiny forest throughout the region from near 
sea level to approximately 1950 m. The only lit- 
toral forest site at which we recorded it was Itap- 
era. Because all of the field work at littoral forest 
sites was during the cuckoo's highly vocal period 
in Madagascar, our failure to find this species 
there indicates that it was absent or rare in these 



forests. The species of birds this cuckoo regularly 
parasitizes, e.g., Terpsiphone mutata and Neomix- 
is tenella (Langrand, 1990), are common in lit- 
toral forest. Cuculus was present in humid forest 
of parcel 1 of the RNI d'Andohahela from 440 m 
to 1950 m, although it was rare above 1500 m. 

Apart from the well-known "tao-tao-kafo" call 
of the male, there is also an infrequently heard 
call, consisting of a loud series of single notes, 
"pee-pee-pee-pee-pee-pee-pee-pee," that is simi- 
lar in tone structure and frequency to calls made 
exclusively by females in other Cuculus spp. 
When this call was replayed in the presence of 
singing males, they typically stopped calling al- 
most immediately and then flew toward the source 
of call. They then started singing again. 

Local Name — Taotaokafo (Berenty, Manom- 
bo, Marosohy). 



Coua gigas 
Giant Coua 

The Giant Coua was a relatively common in- 
habitant of spiny forest and gallery forest, includ- 
ing the transitional forest of parcel 3 of the RNI 
d'Andohahela and the littoral forest as far north 
as Manafiafy. This species was regularly heard 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



45 



and seen in the degraded Mandena Forest but was 
not recorded in the Bezavona Forest, with lateritic 
soils, a few kilometers away. In mid-October the 
Giant Coua was one of the most frequently heard 
birds in the Itapera Forest. Here, as in the re- 
maining littoral forest fragments of Manafiafy and 
Mandena, this species was often found in parcels 
of forest less than a few hectares in size and was 
observed walking about in heavily disturbed areas 
between forest patches. Thus, it appears that this 
species is tolerant of considerable forest degra- 
dation and is able to move between forest frag- 
ments. In the RP de Berenty the densest popula- 
tions occur in the tall closed-canopy gallery for- 
est, where the leaf litter is thickest. This species 
was common in spiny forest adjacent to gallery 
forest along the Mananara River in parcel 2 of the 
RNI d'Andohahela. Bluntschli collected one spec- 
imen near Amboasary-Sud on 22 October 1931 
(smf); at this time extensive gallery forest re- 
mained along the banks of the Mandrare River. 

On 17 December in the Malaza Forest a "large 
boa (Acrantophis dumerilii) [was] seen striking, 
constricting and killing [a] ... Coua gigas; [an 
adjacent] bird went berserk, calling and flapping 
its wings from a perch 2-3 feet above the scene. 
Death was slow and the [bird] . . . continued flap- 
ping . . . [its] wings for minutes after the strike" 
(SOC). 

Given its terrestrial habits, C. gigas is probably 
easy for local people to capture. Indeed, at most 
sites where this species was recorded there was 
direct or indirect evidence that it is hunted for 
food by local people. Hunting techniques included 
slingshots, snares, and walk-in traps. A local vil- 
lager at Ankapoky mentioned that his sons were 
able to capture at least one Giant Coua every 2 
weeks and that this meat was an important sup- 
plement to the family's diet. At Bealoka in early 
1984, one or two pairs were present, but by 1987 
this species was absent. Its disappearance is pre- 
sumably related to hunting. 

Diet — The stomachs of two collected birds 
contained Diplopoda, beetles (Carabidae, Curcu- 
lionidae, Scarabaeidae, Tenebrionidae), ants (For- 
micidae), flies (Asilidae), lepidopteran larva, and 
small seeds (Goodman & Parrillo, in press). The 
stomach of a specimen collected at Mandena on 
28 July contained remains of grasshoppers and 
various other insects (mnhn). 

Foraging often involves chasing down insect or 
small reptilian prey. Giant Couas react and run 
very quickly. A coua in pursuit of prey may 
change directions at very acute angles, often 



pushing off trees or other vertical surfaces with 
the feet and legs. It has been observed leaping a 
meter or so into the air from the ground to catch 
prey in flight. This species also scrapes in leaf 
litter in search of food. 

Breeding — Nests of this species found in the 
RP de Berenty were built by both males and fe- 
males. If birds are discovered during nest building 
they will wait long periods before returning to the 
nest or take alternative routes to keep the location 
secret. The nests are relatively elaborate structures 
composed of large twigs or small branches in 
Acacia or Tamarindus trees and usually in vines 
and lianas in areas of dense vegetation. In the RP 
de Berenty nest building has been observed be- 
tween late October and late December, and chicks 
have been seen in January. Two males collected 
at Petriky in late September had testes 10 X 8 
mm and 10 X 5 mm. 

Weight— Combined (2), 410, 415 g. 

Soft Part Colors — Bill and legs: black; iris: 
deep reddish brown; orbital ring: above eye bright 
greenish blue, below and posterior to eye pur- 
plish, and anterior to eye grayish blue. 

Local Names — Eoka and aoka (Berenty). 



Coua reynaudii 
Red-fronted Coua 

The Red-fronted Coua was widespread in the 
humid forests of the region from the Marovony 
Forest south through the Anosyenne and Vohi- 
mena mountains to the Manantantely Forest, be- 
tween 50 and about 1950 m. This species was 
relatively common in the forests of Marovony, 
Analalava, Bemangidy, Marosohy, Bezavona, and 
Manantantely. In parcel 1 of the RNI d'Andohahela 
it was much less frequently recorded on point 
counts at 440 m and 810 m than at 1200, 1500, 
and 1800 m. It was not recorded at any littoral 
forest site. 

On 26 December 1992 feathers of at least two 
adult Red-fronted Couas were found on the 
ground under a rock overhang along the Isedro 
Trail in parcel 1 of the RNI d'Andohahela at about 
425 m. Also, in early October 1995 feathers of 
this species were found along the Tanatana Trail 
between Isaka-Ivondro and Eminiminy. These 
birds probably were killed by hunters, most likely 
with a slingshot. 

Diet — The stomach of an individual collected 
in the Marovony Forest contained Araneae, vari- 
ous types of Coleoptera (Cerambycidae, Curcu- 



46 



FIELDIANA: ZOOLOGY 



lionidae, Elateridae, Scarabaeidae), Orthoptera 
(Euschmidtiidae, Tetrigidae), and Phasmatodea 
(Goodman & Parrillo, in press). This species will 
use forest clearings and edge habitat along trails 
or roads to search for food among dense vine tan- 
gles. 

Breeding — In parcel 1 of the RNI d'Andohahela 
a presumed female was building a nest at 1200 m 
on 16 November. It picked up dead ferns, bits of 
liana, and roots for construction material. The pre- 
sumed male accompanied the female for a few 
minutes while the female looked for nesting ma- 
terial. The bird then captured a 3-cm-long cater- 
pillar and offered it to the female. They attempted 
to mate, and the male departed while the female 
continued with nest construction. The nest was a 
wide shallow cup built in a tangle of vegetation 
at the end of a fallen tree about 2 m off the 
ground. 

Weight — Sex unknown (1), 163 g. 

Soft Part Colors — Bill: black; legs: slate col- 
ored; claws: black; iris: brown; orbital ring: pos- 
terior to eye bright sky blue, and anterior to eye 
cobalt blue. 

Local Names — Pokafo (Marosohy and parcel 
1, RNI d'Andohahela), taitoaky (Marovony). 



Coua cursor 
Running Coua 

The Running Coua was an inhabitant of the 
spiny forest. The eastern limit of its range appears 
to be the lower western slopes of the Anosyenne 
Mountains. It was recorded at the edge of the tran- 
sitional forest, near Mahamavo, just below the Col 
d'Ambatomaniha on the west side of parcel 1 of 
the RNI d'Andohahela. This species occurs in 
spiny forests but not in gallery forest. It was fairly 
common in spiny forest near the Mananara River 
in parcel 2 of the RNI d'Andohahela, although it 
was the least frequently recorded of the four Coua 
species present at this site. 

Diet — One bird from Ankapoky had spiders 
(Araneae), beetles (Curculionidae), cicadas (Ci- 
cadidae), ants (Formicidae), and plant material in 
its stomach (Goodman & Parrillo, in press). 

Breeding — A female collected at Ankapoky on 
9 October had an enlarged ovary with follicles up 
to 10 mm. 

Weight— Female (1), 118 g. 

Soft Part Colors — Bill, legs, and claws: 
black; iris: brown; orbital ring: deep blue with 
purple tint. 



Local Name — Aliotsy (Berenty). 



Coua ruficeps olivaceiceps 
Red-capped Coua 

The Red-capped Coua was relatively common 
in the spiny forest region, including parcels 2 and 
3 of the RNI d'Andohahela south to at least the 
Bevilany and Fotsivolo hills. Langrand (1990) 
noted that it occurs as far south as Lac Anony. 
Milon collected an adult female, referable to C. r. 
olivaceiceps, near Ranopiso on 31 May (mnhn). 
In December 1995 this species was the most com- 
mon Coua in the spiny forest along the Mananara 
River in parcel 2 of the RNI d'Andohahela. 

Diet — The stomach of the Ranopiso specimen 
contained insects and numerous seeds. 

Breeding — This Ranopiso specimen had an 
ovary 9 X 4.5 mm with no enlarged egg follicles. 
In December east of Hazofotsy, a juvenile was 
entering postjuvenile molt. 

Local Name — Aliotsy (Berenty). 



Coua cristata maxima and Coua cristata pyr- 
opyga 

Crested Coua 

The Crested Coua was an inhabitant of the 
spiny forest region. Its current eastern limit ap- 
pears to be the lower western slopes of the Ano- 
syenne Mountains, where it was recorded near 
Mahamavo, Evasia, and Tanambao-Ankatsaky, 
east along the coast as far as the Petriky Forest, 
and south to the Andraraky Hills due east of Mo- 
kobe. Throughout this region it was relatively 
common and is represented by the subspecies C. 
c. pyropyga. 

In 1950 Milon described a new subspecies of 
Crested Coua, C. c. maxima, from the environs of 
Fort-Dauphin (= Tolagnaro). Stresemann (1952) 
suggested that the type specimen of C. cristata 
also came from the Tolagnaro area (but see be- 
low). The holotype of maxima, the only known 
specimen of this form, is distinctly larger than C. 
c. pyropyga, with no overlap between these forms 
in four external measurements (Table 4). Further, 
maxima differs from typical pyropyga in plumage 
coloration. The following comparisons are based 
on the holotype of C. c. maxima (mnhn 1950.392) 
and a series of C. c. pyropyga from several lo- 
calities in the southwest (mnhn): the upper tail 
surface of maxima is an intense violet blue (rem- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



47 



Table 4. External measurements (in mm) of Coua 
cristata maxima and Coua cristata pyropyga, based on 
Milon (1950). 



Measurement 



maxima pyropyga 

(N =1) (N = 14)' 



Wing 

Tail 

Tarsus 

Bill (from commissure) 



175 162 (157-168) 

232.5 212 (208-224) 

45 41.2(38.5-43.5) 

30 26.7 (25-28.5) 



Mean (range). 



iniscent of C. caerulea) as compared with the 
more subdued iridescent blue with a greenish 
tinge in pyropyga; the innermost secondaries of 
maxima are distinctly blue, whereas in pyropyga 
they are iridescent green; the back of maxima is 
a grayish blue and that of pyropyga is gray or 
greenish gray; in pyropyga the lower breast is 
white with a rufous vent and in maxima it is taw- 
ny brown with no clear sign of a change in col- 
oration at the vent (although the specimen appar- 
ently lost most of the undertail coverts during 
preparation); and in pyropyga the white belly 
merges anteriorly from a light tawny brown lower 
mid-breast to a purplish gray mid-breast and low- 
er throat and ends in a gray throat, whereas in 
maxima the tawny brown lower breast merges to 
a darker tawny brown upper breast and cold blu- 
ish gray throat. Milon (1950) noted that the ven- 
trum of the maxima specimen was tawny cinna- 
mon with no reddish coloration at the base of the 
tail. 

We failed to find anything resembling maxima 
in any area in which we worked in southeastern 
Madagascar. Most of the natural lowland forest in 
the immediate vicinity of Tolagnaro has been de- 
stroyed. One of the last vestiges of somewhat in- 
tact habitat was near Lac Lanirano, 2 km north of 
Tolagnaro, where one C. cristata was observed 
and heard calling in August 1988; by 1989 this 
individual apparently had disappeared. A search 
for this species in the small remnant forest of Lac 
Lanirano in late December 1992 was unsuccess- 
ful. No particular note was made at the time of 
plumage characters of the bird observed at Lac 
Lanirano in 1988, and it is not known whether 
this bird was maxima. This possibility may be un- 
likely, because Milon (1950) noted that in life the 
type of maxima was easily confused at first view 
with C. caerulea. 

Stresemann ( 1 952) reported that the holotype of 
Coua cristata was collected in 1752 by Poivre in 



the Fort-Dauphin area; nothing remains of these 
collections. The Poivre specimen, "Coucou hupe 
de Madagascar," was the holotype of Cuculus 
cristatus L. 1776 and was illustrated by Dauben- 
ton (1770-1786, plate 589). Although somewhat 
stylized, this color plate clearly shows a bird with 
rufous coloration in the undertail coverts and a 
white lower ventrum. Further, Brisson's (1760, p. 
1 50) description of the specimen notes that it has 
a white underside and that the feathers at the base 
of the tail are reddish white (our translation). 
Thus, on the basis of these characters the Poivre 
specimen is not referable to C. c. maxima. Further, 
the Poivre specimen had a reddish venter, which 
opens up the taxonomic problem that the holotype 
of nominate cristata, which is distinguished in 
part from C. c. pyropyga by its tawny undertail 
coverts (Delacour, 1931), probably represents the 
population now known as pyropyga. 

The taxonomic status of maxima is unclear. At 
this stage it could be a geographic form of C. 
cristata, a distinct species, or even a hybrid be- 
tween C. cristata and one of the two other couas 
living in the region (i.e., C. caerulea and C. rey- 
naudii). Current research on the intrageneric re- 
lationships of the couas might help to resolve this 
question. 

The southern limit of C. c. cristata in eastern 
Madagascar appears to be in the vicinity of RS de 
Manombo, south of Farafangana (Nicoll & Lan- 
grand, 1989). This locality is approximately 130 
km north of the Marovony Forest. Thus, this spe- 
cies is absent from the humid and littoral forests 
(excluding Petriky) of southeastern Madagascar. 

The calls of C. cristata are loud and carry for 
long distances. Often the calls of one bird will 
initiate responses from others, especially at dawn 
and dusk. Such a bout of countercalling may con- 
tinue quite far through a gallery forest as 15 or 
more birds respond to one another. 

Diet — Coua cristata gleans prey from trunks 
and branches of large mature trees of Albizia po- 
lyphylla and Acacia rovumae. Starting at the base 
of the tree it progresses systematically up the 
trunk while rapidly checking nooks and crannies 
and under leaves. At the top of the tree it then 
glides to the base of the next tree, starting the 
process over again. It often walks or hops up 
along nonvertical tree trunks and branches, and 
invariably the process includes short bursts of fly- 
ing and gliding. 

Specimens collected near Petriky had Coleop- 
tera (Curculionidae), Hemiptera, Homoptera, lep- 
idopteran larvae, Mantodea, orthopterans (Acrid- 



48 



FIELDIANA: ZOOLOGY 






idae, Euschmidtidae, Gryllidae, Pyrgomorphidae), 
Phasmatodea (Phylliidae), and plant seeds in their 
stomachs (Goodman & Parrillo, in press). Birds 
have been seen carrying lizards (Mabuya), geck- 
os, chameleons, locusts, and beetles in their bills. 
C. cristata is often observed in Albizia polyphylla 
trees searching for prey. This tree also exudes co- 
pious quantities of resin that this coua feeds on. 
In spiny forest this species had been previously 
reported to feed on sap (Charles-Dominique, 
1976). In the humid forest of Manombo we saw 
it feeding on sap. It also has been observed eating 
tree flower buds in the RP de Berenty (OL & 
LW). 

Breeding — In the RP de Berenty young fledg- 
lings have been observed from late October to 
early February. Fledglings beg by fluttering their 
wings while uttering a humming-hissing vocal- 
ization. Several nests were found at the RP de 
Berenty in the understory of Celtis phillipensis 
trees. Three adult males taken at Petriky in late 
September had testes ranging in size from 4x2 
mm to 10 x 4 mm. A female collected at Anka- 
poky on 1 1 October had a vascularized brood 
patch, a shelled egg in the oviduct, ovarian folli- 
cles 26 and 16 mm, and no corpus luteum. On 15 
November in the Hazofotsy Forest an adult was 
observed feeding a fledgling capable of flight, and 
on 8 December at a nearby site a juvenile was 
being attended by two adults. Nests have been 
found in January near Hazofotsy in a Moringa 
tree. 

Weight— Combined (6), 144.2 ± 7.5 (135- 
152) g. 

Soft Part Colors — All pyropyga. Bill and 
legs: black; iris: red, bright purplish blue, or red- 
dish brown; orbital ring: posterior to eye mixed 
sky blue or greenish blue and brilliant green and 
anterior to eye cobalt blue, bright purplish blue, 
or purplish brown. 

Local names — Fandikalala (Manombo), tivo- 
ka (Berenty). 



[Coua verreauxi 
Verreaux's Coua 

This species has been reported from the RP de 
Berenty (Langrand, 1990). The juvenile plumage 
of C. cristata can be easily confused with that of 
C. verreauxi, and we suspect that records of C. 
verreauxi from this locality are in error. Until fur- 
ther documentation is available this species is not 
considered to occur in southeastern Madagascar.] 



Coua caerulea 
Blue Coua 

The Blue Coua was the most common of the 
humid forest couas. It was recorded from the Ma- 
rovony Forest south through the Anosyenne and 
Vohimena mountains to Manantantely and Pic St. 
Louis from near sea level to 1800 m. In parcel 1 
of the RNI d'Andohahela this species was ob- 
served between 440 and 1 800 m but was recorded 
most commonly on point counts at 440, 810, and 
1200 m and was much more scarce at 1500 and 
1800 m. During the 1989 and 1990 field seasons, 
this species was relatively common in the littoral 
forests of Manafiafy and Itapera but was absent 
from Mandena. At Mandena one individual was 
observed in May 1988, and presumably it was 
locally uncommon or perhaps moved seasonally 
between the littoral forest and nearby humid forest 
on lateritic soils. There are several older records 
from the Tolagnaro area (Lavauden, 1937; Milon, 
1952), including a specimen taken in 1756 near 
Fort-Dauphin (Stresemann, 1952). This species 
still occurs at Pic St. Louis, 3 km from Tolagnaro. 
The Blue Coua was not recorded in the remaining 
coastal forest patches west of Tolagnaro (e.g., Pe- 
triky) or in the spiny forest region. Severe hunting 
pressure combined with habitat destruction over 
the past few decades, particularly in forests near 
villages, may have greatly altered the distribution 
of this species. 

Nowhere in southeastern Madagascar is the 
Blue Coua sympatric with the Crested Coua, and 
in this region these two species of arboreal couas 
appear to replace one another geographically. In 
the humid forest below the Col d'Ambatomaniha, 
on the western side of parcel 1 of the RNI 
d'Andohahela, the Blue Coua was relatively com- 
mon. At about 810 m, when the abrupt transition 
to dry forest commences, this species dropped 
out, and by 650 m the Crested Coua was common. 
This change is over a ground distance of less than 
2 km. To the north, as in the RS de Manombo 
( 1 30 km north of the Marovony Forest), these two 
species are sympatric in humid forest. 

Diet — On several occasions we observed the 
Blue Coua in littoral forest feeding on a resin-like 
substance exuded from the ripe fruits and trunk 
of Sloanea rhodantha. The stomachs and lower 
intestines of two individuals taken at Manafiafy 
and of one taken at Marovony contained a sticky, 
viscous, and non-water-soluble liquid. Stomach 
contents of individuals collected in the general 
Tolagnaro area (fmnh, mnhn) included various 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



49 



types of insects (often grasshoppers) and a large 
millipede 90 X 7 mm. One bird taken in littoral 
forest had a centipede (Scolopendromorpha) and 
a Phasmatodea in its stomach (Goodman & Par- 
rillo, in press). On several occasions we observed 
this species feeding on insects either on the 
ground or at various levels in the forest understo- 
ry. 

In humid forest at 1500 m in the RNI 
d'Andohahela, an adult was observed holding a 
Calumma nasutus chameleon and a large katydid 
in its bill, which were fed to a second Blue Coua; 
this was probably an example of courtship feed- 
ing. In the Bezavona Forest a Blue Coua was not- 
ed feeding on the unopen flower buds of Sym- 
phonia. 

On several occasions we observed C. caerulea 
following within a few meters behind troops (four 
to over 15 individuals) of the Brown Lemur (Eu- 
lemur fulvus). This association was noted in both 
littoral and lowland humid forests and always in- 
volved a single bird. E. fulvus generally moves in 
the lower to middle portion of the forest and is 
largely folivorous and frugivorous (Richard & 
Dewar, 1991); this species also has been recorded 
eating invertebrates (spiders, millipedes, cicadas) 
and fungi and, at least in captivity, vertebrates 
(Glander et al., 1985; O'Connor, 1987). Although 
no prey was caught by the Blue Couas that we 
observed following troops, it is likely that the 
movements of the lemurs would scare or expose 
prey upon which the bird may feed. 

Breeding — There was variation in the breeding 
condition of Blue Couas obtained at Manafiafy in 
October. A male in adult plumage had testes 3 X 
1 mm, whereas the testes of two other adult males 
were (left and right, respectively) 12X5 mm and 
9X4 mm, and 15 X 6 mm and 10 X 5 mm. An 
adult female taken during the same period had 
ovarian follicles 16 X 14 mm, 11X11 mm, and 
4X4 mm and had one ruptured follicle. A male 
collected at Marovony on 2 November had testes 
13 X 9 mm (left) and 12 X 7 mm (right). 

Milon collected two specimens on 28 May 30 
km NNW Tolagnaro: a female with a slightly en- 
larged ovary and an ovarian follicle of 9 mm and 
a male with testes 3.5 X 1 mm (left) and 4 X 1 
mm (right) (mnhn). A female taken 7 km north of 
Tolagnaro had a nonenlarged ovary (mnhn). 

On 15 October in the littoral forest of Manafia- 
fy a Blue Coua was heard giving a "purr-like" 
call from a 1.5-m-high perch. The coua held a 
frog (Heterixalus boettgeri) in its bill. The frog 
was positioned in the bird's beak so that its bright 



yellowish orange legs and ventrum were conspic- 
uously exposed. The coua gave a "purr-call" as 
it slowly lowered and raised its head every 2-3 
seconds. This sequences was repeated numerous 
times. After every two or three such sequences 
the bird would flick the wings open to a one-quar- 
ter to one-half position, immediately close them, 
and then lift the tail, open it into a fan-like posi- 
tion, and then close it. After about 10 minutes of 
repeating this display another Blue Coua called 
from about 40 m away. At this point the intensity 
of the display increased at least twofold, and the 
displaying bird completely opened the wings and 
fanned the tail with each bow. After the second 
bird flew off, the intensity and frequency of the 
display returned to the original level. The bird 
then was disturbed by someone passing by in the 
forest and flew off. 

Weight— Combined (6), 235.3 ±11.7 (225- 
257) g. 

Soft Part Colors — Bill, legs, and claws: 
black; iris: brown or reddish brown; orbital ring: 
anterior to eye purplish blue and posterior to eye 
cobalt blue or in general orbital ring cobalt blue 
or violet blue. 

Local Names — Teso (parcel 1 of the RNI 
d'Andohahela), tetso (Manombo, Marosohy), tet- 
so manga (Manafiafy), tivoka (Manafiafy). 



Centropus toulou toulou 
Madagascar Coucal 

The Madagascar Coucal inhabits a variety of 
habitats. It was most common in thick secondary 
scrub and other types of dense vegetation, al- 
though it can be found in relatively intact littoral, 
humid, spiny, and gallery forests. It also occurs 
in Phragmites communis marshes, thick grass- 
lands, vegetated fence rows at the edge of villages 
and gardens, and eucalyptus and sisal plantations. 
Its elevational range in the region was from near 
sea level to about 1700 m. In the RNI 
d'Andohahela (parcel 1) this species was relative- 
ly scarce and was recorded at 440, 1200, and 1500 
m in intact humid forest and at 1700 m in tran- 
sitional forest north of Trafonaomby. 

Female C. toulou are significantly larger than 
males (Andersson, 1995; our weight data below), 
as is the case for a number of other species in this 
genus (Andersson, 1995). An African species of 
Centropus, C. grillii, which is thought to be close- 
ly related to C. toulou (Snow, 1978), is polyan- 
drous (Vernon, 1971), as might be expected from 



50 



FIELDIANA: ZOOLOGY 



a species that shows reversed size dimorphism be- 
tween the sexes (Andersson, 1995). The mating 
system of C. toulou is not known but is worthy 
of study. We strongly suspect that polyandry will 
be demonstrated in this species as well. 

Diet — The stomach of a specimen taken near 
Mandena (mnhn) contained debris of spiders and 
grasshoppers. Birds collected in littoral forest ar- 
eas had a wide assortment of remains in their giz- 
zards, including centipedes (Scolopendromorpha), 
spiders (Araneae, Salticidae), cockroaches (Blat- 
todea), beetles (Elateridae, Scarabaeidae, Teneb- 
rionidae, Tettigoniidae), ants (Formicidae), crick- 
ets (Gryllacrididae), mantids (Mantidae), and a 
small gecko (Goodman & Parrillo. in press). 
Adults were observed on 12 December carrying 
a large grasshopper and a smaller cricket to a nest 
placed in riverside vegetation along the Mananara 
River. 

Breeding — Four specimens taken near Mana- 
fiafy in the second half of October were in or 
approaching breeding condition. These included 
two males (both with degenerated left testes); one 
had a 6-mm bursa and the right testicle was 12 X 
9 mm, and the other had no bursa and the right 
testicle was 15 x 7 mm. Rand (1933) also noted 
testicular asymmetry in this species. A female 
taken on 16 October had a shelled egg in the ova- 
ry and two corpora lutea, and another obtained on 
23 October had a thickened oviduct, no brood 
patch, and three corpora lutea. A female taken at 
Mandena on 16 September had a 30 x 20 mm 
unshelled egg in the oviduct and one ruptured 
ovarian follicle; the largest intact follicles were 10 
and 7 mm. 

Weight— Female (1), 220 g; male (4), 150, 
180, 185, 210 g. 

Soft Part Colors — Bill: generally black, al- 
though in one adult male the mandible tip was 
gray; legs: dull or dark bluish gray to black; iris: 
deep red. 

Local Names — Kotohake (Berenty), toloho 
(Berenty, Manafiafy, Manombo, Marosohy). 



Strigiformes 

Tytonidae 

Tyto alba affinis 
Barn Owl 

We have few records of the Barn Owl. Presum- 
ably it occurs throughout the region, particularly 



in agricultural areas and as a human commensal 
(Goodman & Langrand, 1993). This species was 
heard calling at night in October and November 
in Tolagnaro proper and in September at Man- 
dena. One Barn Owl was found dead along Route 
Nationale 13, 14 km west of Tolagnaro in an area 
of open agricultural land and eucalyptus planta- 
tion. This species occurs in gallery forest along 
the Mandrare River, particularly in the Malaza 
Forest, and in open areas outside of the forest. 

Diet — The bird found dead along the road had 
an adult Rattus in its stomach. A small collection 
of pellets picked up in the Kaleta Reserve along 
the Mandrare River predominantly contained Rat- 
tus rattus and a few individuals of Geogale aurita 
and Microcebus murinus. 

Local Names — Haka (Manombo), heko heko 
(Berenty), hora (Marosohy), vorondolo (Berenty, 
Manafiafy). 



Strigidae 

Otus rutilus rutilus 
Malagasy Scops Owl 

The Malagasy Scops Owl was a common in- 
habitant from the Marovony Forest south through 
the Anosyenne and Vohimena mountains and 
along the coast to Petriky and west to the Man- 
drare River. This species occurs in a wide variety 
of habitats, including intact to disturbed humid, 
littoral, transitional, spiny, and gallery forests be- 
tween near sea level and 1500 m. It is broadly 
sympatric with Ninox throughout the spiny forest 
region and locally in littoral forest at Petriky. This 
species is known from the Amboasary-Sud area 
(Langrand & Meyburg, 1984) and the RP de Ber- 
enty. 

Two call types of this species have been re- 
ported from Madagascar. Rand ( 1 936, p. 39 1 ) de- 
scribed the song as "tura-tura-tura-tura ,*' i.e., 

a series of two-parted calls. Langrand (1990, p. 
227), however, presented the song as "hoo hoo 
hoo hoo hoo hoo hoo or broo broo broo, 5-7 
notes delivered at same pitch." The "hoo" se- 
quence clearly is a series of a single note, al- 
though "broo" may implicitly describe a two-not- 
ed call. Marshall (1978, plate 10) provides sona- 
grams of the two song types. 

Marshall (1978, p. 19) reported that Stuart 
Keith and the late C. W Benson had observed that 
the voice of Otus rutilus was "variable" and, 
without elaboration, that "Keith heard them [i.e., 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



51 



the two song types] in different places." Our ex- 
perience also has been that songs of Otus rutilus 
are geographically variable. The lower, slightly 
rougher, two-parted call is typical of birds we 
have heard in the RP de Berenty and parcel 2 of 
the RNI d'Andohahela and at other sites in west- 
ern Madagascar (e.g., Kirindy Forest, RNI de Na- 
moroka, RNI d'Ankarafantsika) in deciduous or 
spiny forest. The higher, smoother, single-note se- 
ries, composed of 8-30 notes, is typical of birds 
of humid forests in southeastern Madagascar, as in 
parcel 1 of the RNI d'Andohahela, Mandena, An- 
alalava, and Marovony and at localities farther north 
(e.g., PN de Ranomafana, RS d'Analamazaotra, 
RNI de Zahamena, RS d'Anjanaharibe-Sud, RNI 
de Marojejy). Marshall (1978) reported that Keith 
had heard one bird "switch" from one song type 
to another. In fact, Keith (in litt., 1990) heard "... 
both song types from the same patch of forest [at 
Fampanambo, near Maroantsetra]." 

Such a pattern of geographic variation in song 
presumably would have a genetic basis. Only the 
nominate form of Otus rutilus is recognized on 
Madagascar, and we also have not detected any 
morphological or plumage variation that might 
correlate with voice. Further analyses of song 
variation in this species are planned to confirm the 
distinctiveness of the two vocal types, especially 
in ecotonal areas. 

Diet — A single specimen taken in the Manan- 
tantely Forest had spiders (Araneae), Chilopoda, 
beetles (Curculionidae, Elateridae, Scarabaeidae), 
and Orthoptera (Gryllidae) in its stomach (Good- 
man & Parrillo, in press). In the Malaza Forest 
this owl fed on cicada nymphs. In the humid for- 
est of the RNI d'Andohahela, this species was ob- 
served feeding on a Calumma nasutus chameleon 
and a large katydid. 

Breeding — Specimens taken between mid-Sep- 
tember and late November had enlarged repro- 
ductive organs. 

Weight— Female (1), 112 g; male (3), 85, 97, 
105 g; unknown (2), 94, 116 g. 

Soft Part Colors — Bill: maxilla generally 
black but in one individual olive gray with dusky 
tip, and mandible black, black with yellowish tip, 
olive with whitish gray tip, or dull beige with 
black cutting edge; cere: olive gray or dull pink; 
legs: light or dull pink; claws: black or gray with 
black tips; iris: ochre yellow or lime yellow; or- 
bital ring: reddish pink. 

Local Names — Toro or torotoroka (Berenty, 
Marosohy). 



Ninox superciliaris 
White-browed Owl 

The White-browed Owl was an inhabitant of 
dry areas, including spiny, gallery, and transition- 
al forest, west of the lower western slopes of the 
Anosyenne Mountains. The only locality from 
which it was recorded outside of this range was 
the coastal forest of Petriky, which appears to be 
the eastern limit of this species' range in southern 
Madagascar. It is unknown from other littoral or 
any humid forests of the region, although in 
northern portions of the island it occurs in humid 
and littoral forest (Langrand, 1990; pers. obs.). 
Bluntschli collected two adult females on 1 1 and 
1 8 November at Amboasary-Sud (amnh, smf), the 
same locality where Langrand and Meyburg 
(1984) found this species. It occasionally can be 
heard calling 2 hours before dusk and after dawn 
and on overcast days during the mid-morning or 
early afternoon. 

This species is sympatric with Otus rutilus in 
spiny and locally in gallery forest. At dusk it vo- 
calizes almost without exception before O. rutilus. 
During the day Ninox often roost in pairs and will 
perch on branches in the dense and darkest part 
of the forest. 

Breeding — Two eggs of this species were col- 
lected from a nest at Amboasary-Sud in mid-No- 
vember (amnh). At the RP de Berenty on 6 No- 
vember a pair was observed, and one bird ap- 
peared to be incubating a nest in a hollow of a 
dead tree (OL). 

Local Name — Vorondolo (Berenty). 



Asio madagascariensis 

Madagascar Long-eared Owl 

The Madagascar Long-eared Owl was recorded 
at several localities from the Marovony Forest 
south to the general Tolagnaro region. This spe- 
cies also has been reported from the RP de Ber- 
enty (Langrand, 1990). It occurs in a variety of 
habitats, including littoral, humid, and gallery for- 
ests from near sea level to about 1 200 m. We have 
recorded this species at three locations in the gen- 
eral Tolagnaro area: in the pristine humid forest 
of Marosohy at about 350 m, in a large and rel- 
atively undisturbed patch of littoral forest at Man- 
afiafy, and in a disturbed area at about 75 m in 
the Bezavona Forest. GR reported this species at 
Mandena. An undated specimen (bmnh) was col- 
lected near Fort-Dauphin sometime before the 



52 



FIELDIANA: ZOOLOGY 



turn of the 20th century. On 14 October Bluntsch- 
li collected one individual near Eminiminy (smf). 

In the humid forest of the RNI d'Andohahela 
(parcel 1) it was heard calling at 440, 810, and 
1200 m. The typical call is a series of three to 10 
"hangh" notes. One individual called on several 
days numerous times between dawn and 12h00 
from a large Canarium tree with dense foliage; 
the call of this bird was a single "hangh." In late 
November and early December Madagascar 
Long-eared Owls were heard calling in the Ma- 
rosohy Forest and in November were heard in the 
Marovony Forest. 

Diet— Pellets collected at a roost site of this 
species in the Bezavona Forest contained amphib- 
ians (Boophis madagascariensis, Platypelis gran- 
dis), reptiles (Uroplatus sikorae), birds (Centro- 
pus, Otus, Eurystomus, Hypsipetes), bats (Hippos- 
ideros commersoni), lemurs (Microcebus, Avahil 
Hapalemur), and rodents (Eliurus minor, E. 
xvebbi, Rattus, Mus) (Goodman et al., 1991 
1993). 

Breeding — In late December a juvenile bird 
was observed in the Bezavona Forest near a roost 
site. 

Local Name — Hankana (Marosohy). 



Asio capensis [hova] 
Marsh Owl 

The Marsh Owl has been observed on a few 
occasions. One was noted resting on a grass tus- 
sock on the shore of Lac Andriamibe, 5 km west 
of Tolagnaro. On several occasions this species 
was observed at night near Lanirano, 2 km north 
of Tolagnaro, perched on a fence around an illu- 
minated compound. This species also has been 
observed in the RP de Berenty (OL; Nagata et al., 
1992). 



Caprimulgiformes 

Caprimulgidae 

Caprimulgus madagascariensis 
madagascariensis 
Madagascar Nightjar 

The Madagascar Nightjar was a common in- 
habitant of disturbed habitats along the coastal 
plain, particularly at forest edge, in open grass- 



lands, agricultural areas, and in towns and villages 
from sea level to 850 m. The highest elevation at 
which we recorded this species was near Antseva, 
in an area of open grassland on the slopes of the 
Anosyenne Mountains. It also was a common in- 
habitant of open areas of spiny forest and along 
gallery forest. We did not record it in intact humid 
forests, such as those of the RNI d'Andohahela. 
Virtually all of the Madagascar Nightjars netted 
during our field studies (N = 14) were captured 
in open areas away from primary or tall second- 
ary forest. The only exception was at Ankapoky, 
where they were captured in relatively open, in- 
tact spiny forest. 

The vocalization of this species is one of the 
characteristic night sounds of the nonforested por- 
tions of the region, often starting before sunset 
and continuing to dawn. At some localities it was 
common; for example, in the early evening of 10 
October near Manafiafy we found six individuals 
calling along a 0.5-km section of road. This spe- 
cies was common at night on roads in the Man- 
drare River Valley and also along the Route Na- 
tional 13 between Amboasary-Sud and Tolag- 
naro. The dirt and tarmac roads are warm in the 
early evening because of absorption of heat from 
the sun during the day. 

A single specimen from Tolagnaro (fmnh) is 
distinctly lighter in plumage coloration than typ- 
ical C. m. madagascariensis. This variation sug- 
gests that further investigation of the taxonomic 
status of the southeastern Madagascar population 
is warranted. 

Diet — Individuals collected during the 1989 
and 1990 field seasons had beetles (Cerambyci- 
dae, Chrysomelidae: Alticinae, Curculionidae, 
Elateridae, Scarabaeidae: Hopliini), Odonata, and 
plant seeds in their stomachs (Goodman & Par- 
rillo, in press). Specimens taken by Milon also 
had insect debris in their stomachs (mnhn). 

Breeding — Bluntschli collected a nestling on 
12 November near Amboasary-Sud (amnh). On 
24 September a juvenile was found on open 
ground at Petriky. Adult males taken between 
mid-September and late October generally had 
testes up to 9 X 4 mm. A female collected on 9 
October at Ankapoky had a shelled egg in the 
oviduct, one corpus luteum, and a 14-mm ovarian 
follicle. A nestling, almost capable of sustained 
flight, was found at the edge of the forest in parcel 
2 of the RNI d'Andohahela on 9 December. 

Weight— Female (4), 47.8 ± 2.5 (45-51) g; 
male (8), 39.8 ± 2.2 (37-43) g; combined (16), 
41.9 ± 4.4(36-51) g. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



53 



Soft Part Colors — Bill: maxilla black, and 
mandible black or pinkish brown with gray or 
black tip; legs: brown; claws: black; iris: dark 
brown. 

Local Names — Kopaky (Manombo), langopa- 
ka (Berenty), langopaka tataro (Manafiafy). 



Caprimulgus enarratus 
Collared Nightjar 

There are few records of the Collared Nightjar. 
One was observed at 700 m in the humid forest 
of parcel 1 of the RNI d'Andohahela roosting dur- 
ing the day under the protection of a concave por- 
tion of a rocky outcrop. Bluntschli collected two 
individuals near Eminiminy in mid-October at ap- 
proximately 300 m (amnh, smf). On 2 October an 
adult female was netted in the Marovony Forest 
in an area of intact forest (fmnh). The net was 
placed perpendicular to a trail in the forest and at 
least 700 m from the edge. On 26 December 1992 
feathers of this species were found on the ground 
under a rock overhang along the Isedro Trail, par- 
cel 1 of the RNI d'Andohahela, at 425 m. The 
bird was presumably killed by a human hunter. 
Milon et al. (1973) reported this species from the 
Tolagnaro area, but specific details are lacking. 

Diet — The stomach of one collected individual 
contained beetles (Scarabaeidae: Melolonthinae 
and Tenebrionidae) (Goodman & Parrillo, in 
press). 

Breeding — The Marovony Forest specimen 
had a slightly enlarged ovary and thickened ovi- 
duct; the largest ovarian follicle was 2 mm. 

Weight — Female (1), 54 g. 

Soft Part Colors — Bill: black; legs: dark 
brown; claws: black; iris: brown. 

Local Names — Goapakala (Marovony), tatar- 
ao-ala (parcel 1, RNI d'Andohahela). 



Apodiformes 

Apodidae 

[Collocalia francica 
Mascarene Swiftlet 

Milne Edwards and Grandidier (1879) men- 
tioned that cliffs near Tolagnaro were an ideal site 
for the Mascarene Swiftlet to nest, but they pro- 
vided no information that this species actually 



bred in the area. Their general statement was ac- 
cepted as evidence of this species occurring on 
Madagascar (Berlioz, 1946). This notion has been 
perpetuated in the literature. In the absence of any 
confirmed records, however, this species has been 
removed from the Madagascar bird list (Langrand 
& Sinclair, 1994).] 



Zoonavena grandidieri [grandidieri] 
Malagasy Spine-tailed Swift 

The vast majority of our records of the Mala- 
gasy Spine-tailed Swift are from the edge of or 
above relatively dense closed-canopy humid for- 
est between 50 and 1950 m. Localities include the 
forests of Analalava, Bezavona, and Marosohy 
and several sites in or near parcel 1 of the RNI 
d'Andohahela. We did not observe this species in 
littoral forest or in open disturbed lowland areas. 
It was occasionally noted in the spiny forest. The 
majority of records from this habitat are from 
along gallery forest, e.g., along the Mananara 
River near Hazofotsy and along the Mandrare 
River near the RP de Berenty. At the former lo- 
cality this species was observed drinking on the 
wing from the river. 

On several occasions this species was noted 
with other swifts. For example, on 9 December, 
at about 375 m in the Marosohy Forest, two Zoon- 
avena were observed screening insects with three 
Apus barbatus and one Cypsiurus parvus. 

Breeding — Although no nest of Zoonavena 
was found in southeastern Madagascar, on several 
occasions it was observed flying around baobabs 
(Adansonia) in parcel 2 of the RNI d'Andohahela. 

Local Names — Fililiotra (Marosohy), tsilotsi- 
lon'aka (Manafiafy); both generic for swallows 
and swifts. 



Cypsiurus parvus [gracilis] 
African Palm Swift 

The African Palm Swift was broadly distributed 
throughout open lowland areas from Marovony 
south to Petriky and west into the spiny forest to 
the Mandrare River. It is one of the characteristic 
lowland species of the region. The elevational 
range of this species was from near sea level to 
about 1200 m. In the RNI d'Andohahela (parcel 
1 ) it was recorded flying over rivers and over the 
forest canopy at 440, 810, and 1200 m, although 
in such habitat it was much rarer than Zoonavena 



54 



FIELDIANA: ZOOLOGY 



grandidieri. In parcel 2 of the same reserve, this 
species was distinctly more common over spiny 
and gallery forest. It was often observed screen- 
ing insects over wetlands such as marshes and rice 
paddies, particularly in areas where scattered 
palms occurred, and in and around villages and 
towns. On numerous occasions it was noted 
screening insects in mixed-species flocks com- 
posed of other swift and swallow species. For ex- 
ample, on 26 December in open grassland areas 
at 150 m at the edge of the RNI d'Andohahela, 
near Eminiminy, we observed a mixed flock of 
five Cypsiurus, five Apus barbatus, and 10 Phed- 
ina borbonica. 

Breeding — On several occasions flocks of Cyp- 
siurus were seen flying around palms (Dypsis de- 
caryi) in parcel 3 of the RNI d'Andohahela (OL). 



Apus melba [wills i] 
Alpine Swift 

The Alpine Swift was observed at several wide- 
ly separated localities. All of our records are from 
relatively open areas or at the forest edge, for ex- 
ample near or above the forests of Marovony, An- 
alalava, Marosohy, Manafiafy, Mandena, Petriky, 
and Ankapoky. Milon et al. (1973) remarked that 
this species bred in the mountains north of Tolag- 
naro. Although we have no direct evidence to sup- 
port this statement, we did receive a report that a 
colony of swift-like birds occurred along a rocky 
outcrop high in the Vohimena Mountains. 

We often observed this species in mixed-spe- 
cies flocks. On 9 October two A. melba were not- 
ed with several Phedina in the spiny forest near 
Ankapoky; on 3 November four A. melba were 
foraging with four A. barbatus at the edge of the 
Marovony Forest; and on 1 1 November a mixed 
flock of A. melba, A. barbatus, and Phedina were 
observed near Analalava. 

In the RNI d'Andohahela mixed flocks of Apus 
swifts were observed regularly over parcels 1 and 
2 between October and December. Flocks gener- 
ally contained 2-10 individuals. One group of 30 
A. melba was seen flying over the summit of Tra- 
fonaomby in early December; most of the birds 
were in active wing molt, suggesting that they 
were adults that had already finished breeding. 
Over parcel 2, a flock of around 100 swifts was 
noted in mid-December, including approximately 
equal numbers of A. melba and A. barbatus. 



Apus barbatus [balstoni] 
African Black Swift 

The African Black Swift was observed on sev- 
eral occasions in various portions of the area. It 
was recorded near the forests of Marovony, Ma- 
rosohy, and Petriky and near the villages and 
towns of Manantenina, Bemangidy, Tolagnaro, 
and Ambovombe. We found no evidence of it 
breeding in this region, and many of our records 
may be of migrant flocks. It was noted several 
times in flocks with Zoonavena, A. melba, Cyp- 
siurus, and Phedina (see those species accounts 
for specific records). 



Coraciiformes 

Alcedinidae 

Alcedo vintsioides vintsioides 
Malagasy Kingfisher 

The Malagasy Kingfisher was commonly found 
near water in forest and open areas from the Ma- 
rovony Forest south through the Anosyenne and 
Vohimena mountains to Petriky and west to the 
Mandrare River, from near sea level to about 850 
m. On the lower slopes and east of the Anosyenne 
Mountains it was found in a variety of habitats 
from open water (fresh and brackish) such as 
lakes, coastal lagoons, and rice paddies to streams 
and wet areas in humid and littoral forest and in 
open grassland away from water. In this region 
we only have a few records of this species away 
from water. In the RNI d'Andohahela (parcel 1) 
it was seen frequently on rivers in primary forest 
up to 850 m. In spiny forest areas it generally is 
restricted to gallery forest and river margins. This 
species is at least occasionally crepuscular; we 
netted one individual about 1 hour before sunrise. 

Diet — The stomachs of all specimens con- 
tained insects, and in one case they were identi- 
fied as orthopterans. 

Breeding — Birds collected between mid-Sep- 
tember and late November included males with 
testes up to 5 X 3 mm and females with ovarian 
follicles 4 mm in diameter. 

Weight— Female (5), 18.1 ± 2.8 (14.5-21.5) 
g; male (3), 15.3 ± 1.0 (14.5-16.5) g; combined 
(15), 16.9 ± 2.5 (13.0-21.5) g. 

Soft Part Colors — Bill: black or brown and 



l GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



55 



sometimes orange or reddish at base; legs and 
claws: orange or reddish orange; iris: dark brown. 
Local Names — Bintitra (Berenty), revitsy 
(Manafiafy, Manombo, Marosohy), vintsy (Man- 
afiafy). 



Ispidina madagascariensis madagascariensis 
Madagascar Pygmy Kingfisher 

The Madagascar Pygmy Kingfisher was a rel- 
atively common inhabitant of a variety of forest 
types from the Marovony Forest south through the 
Anosyenne and Vohimena mountains to the To- 
lagnaro region and west through scattered loca- 
tions in the spiny forest region to the Mandrare 
River. Its elevational range was from 20 to 1200 m. 
This species was particularly common in pristine 
and relatively intact humid forests such as Ma- 
rovony, Marosohy, Bezavona, RNI d'Andohahela 
(parcel 1 ), and Manantantely and to a lesser extent 
in littoral and spiny forest. One was collected by 
Bluntschli at Eminiminy on 28 October (smf). 
Spiny forest sites include the Ankapoky Forest, 
near Hazofotsy, and near Amboasary-Sud (Lan- 
grand, 1990). A single individual was present in 
the RP de Berenty for three days in May 1986, 
after which it disappeared. 

This species was observed crossing open areas 
such as rice paddies, agricultural fields surround- 
ed by thick secondary forest, and eucalyptus plan- 
tations. In the Bezavona Forest several birds were 
netted along a stream coursing through bamboo 
and eucalyptus at least 500 m from the forest 
edge. 

Diet — The stomach of a collected bird con- 
tained ants (Formicidae) and remains of a small 
reptile (Goodman & Parrillo, in press). One spec- 
imen collected in littoral forest at Mandena had 
bones of a small frog in its stomach. In the Ma- 
rosohy Forest at 750 m this species was observed 
taking stream-dwelling Mantidactylus lugubris. 
Two of these frogs appeared to be chasing one 
another across open rocks along a stream when a 
kingfisher caught one of the frogs. At the same 
locality a collected bird had frog bones in its 
stomach. At 850 m in parcel 1 of the RNI 
d'Andohahela, an Ispidina caught and ate a cha- 
meleon (Calumma nasutus). The body of the cha- 
meleon was about the same length as the king- 
fisher's head. The chameleon was dispatched by 
rapid flicks of its head against a branch. The bird 
kept the prey in its bill for several minutes after 
capture while calling quietly. The kingfisher then 



swallowed the chameleon whole, head first, in 
about 20 seconds. In the same reserve, at 440 m, 
an Ispidina was seen hunting before sunrise in 
near total darkness. 

Breeding — There was considerable variation in 
the development of the reproductive organs of 
adults collected between mid-September and late 
December, from enlarged to small. 

Weight— Female (3), 15.5, 16.5, 22 g; male 
(3), 15, 19, 23.5 g; combined (16), 18.9 ± 2.5 
(15.0-23.5) g. 

Soft Part Colors — Bill: generally completely 
orange or reddish orange but a few individuals 
with dusky brown on central portion of maxilla; 
legs and claws: orange or reddish orange; iris: 
brown. 

Local Name — Revitsy' ala (Manafiafy, Maro- 
sohy). 



Meropidae 

Merops superciliosus superciliosus 
Madagascar Bee-eater 

The Madagascar Bee-eater was relatively com- 
mon in open and heavily disturbed areas, partic- 
ularly along the coastal plain and river margins 
between near sea level and 750 m. On the lower 
slopes and east of the Anosyenne Mountains it 
commonly was seen in gardens of villages and 
towns, at the edge of degraded humid and littoral 
forests, and in agricultural fields or grasslands and 
occasionally flying over relatively intact forest 
parcels. This species was not recorded in 7 weeks 
of bird survey work in parcel 1 of the RNI 
d'Andohahela (October-December), despite 23.5 
hours of canopy watching. It was observed, how- 
ever, at about 1000 m in degraded transitional for- 
est between Trafonaomby and Esomony. In the 
spiny forest region it was noted in a variety of 
habitats, including in intact and degraded spiny 
forest, along riverbanks, at the edge of gallery for- 
est, in eucalyptus and sisal plantations, and along 
roads. In the Bealoka and Malaza forests this spe- 
cies was rare or absent between the months of 
December and February. 

Diet — The stomach of a collected bird con- 
tained Hymenoptera (Apoidea) (Goodman & Par- 
rillo, in press). 

Breeding — Birds collected between late Sep- 
tember and early November included a male with 
testes 6X4 mm and a female with a thickened 
oviduct and ovarian follicles 4 mm in diameter. 



56 



FIELDIANA: ZOOLOGY 



Pairs were observed copulating in Tolagnaro on 
15 October and in the Malaza Forest on 10 Oc- 
tober. Nest holes were relatively common in the 
banks of the Mandrare River between Berenty and 
Bealoka, along the Mananara River, and along the 
Route Nationale 13 between Amboasary-Sud and 
Tolagnaro. 

Weight— Combined (3), 39.5, 45, 48 g. 

Soft Part Colors — Bill: black; legs: grayish 
brown or slate gray; iris: dark brown. 

Local Names — Kirikirioky (Berenty, Manafia- 
fy, Marosohy), kirio (Manombo), tsikiriokirioke 
(Berenty). 



large numbers and tends to take over nest sites in 
tree hollows and cavities that may have recently 
been used by Coracopsis or Agapornis. Eurysto- 
mus is aggressive and highly vocal during the 
breeding season. Favorite nesting trees are dead 
Acacia rovumae, Neotina isoneura, and Tamar- 
indus indica. 

Weight— Female (1), 180 g. 

Soft Part Colors — Bill: yellow with slight or- 
ange cast; legs: brownish olive; claws: black; iris: 
brown. 

Local Names — Tsiraraka (Berenty), teraratsy 
(Marovony). 



Coraciidae 

Eurystomus glaucurus glaucurus 
Broad-billed Roller 

The Broad-billed Roller migrates to east Africa 
during the austral winter. Our earliest record of it 
in southeastern Madagascar is 30 September, and 
the latest record is 2 March. It was a relatively 
common bird throughout the area from near sea 
level to 1600 m. In humid portions of the region, 
from the Anosyenne Mountains east to the sea 
coast, this species occurs in open habitats, includ- 
ing patches of raw deep in the forest, open grass- 
lands, agricultural areas, tree plantations, gardens 
in villages and towns, and open secondary forest. 
In intact humid forest it is rare or absent. In dry 
areas it is a relatively common resident of spiny 
forest and gallery forest and open degraded areas. 
Several specimens were collected at Amboasary- 
Sud in late October and early November (amnh, 
smf). 

Diet — The stomach contents of one collected 
individual contained remains of Elateridae and 
Scarabaeidae beetles (Goodman & Parrillo, in 
press). Like Falco concolor, Eurystomus arrives 
in southeastern Madagascar during a period that 
coincides with cicada emergence, on which they 
feed extensively. This species has been observed 
skimming the water of the Mandrare River; it was 
uncertain whether the bird was drinking or hunt- 
ing. 

Breeding — A female obtained on 2 November 
at the edge of the Marovony Forest had an en- 
larged ovary and thickened oviduct, and the larg- 
est ovarian follicle was 7 mm. The bird did not 
have a brood patch. Active nests have been found 
from early November to early February. 

At the RP de Berenty this species arrives in 



Brachypteraciidae 

Brachypteracias leptosomus 
Short-legged Ground-Roller 

The Short-legged Ground-Roller was relatively 
common in the intact portions of the Marosohy 
Forest between 700 and 1 100 m. It was generally 
noted in areas with large trees, wet ground, and a 
relatively open understory. Although no nests 
were found, birds were frequently flushed up from 
the ground next to hollows and crevices under tree 
buttresses and roots. At least 1 1 birds were ob- 
served along a portion of the Entseva-Anakara 
trail about 1.5 km long between 750 and 825 m. 
Virtually each day during 1 week in late Novem- 
ber individuals were found in the same places 
along this trail. Thus, our impression was that 
these were birds on territory. If so, this would 
give a density of approximately 3.5 pairs/km of 
trail. In the Marosohy Forest this species appears 
to occupy a higher elevational zone than B. squa- 
miger. In parcel 1 of the RNI d'Andohahela, B. 
leptosomus was observed at 440, 810, and 1200 
m, and the local density was not as high as in the 
Marosohy Forest. In early October 1995 feathers 
of this species were found along the Tanatana 
Trail between Isaka-Ivondro and Eminiminy. 
These remains were presumably of birds taken by 
hunters using slingshots. 

The song of this species is a distinctive deep, 
low, and whistled "whop" repeated every 1-2 
seconds. The song is given from a horizontal 
perch 5-30 m high. Individuals regularly can be 
heard countersinging, and apparently territorial 
birds on adjacent territories may chase each other 
between song bouts. 

Diet — One individual was captured after it was 
attracted to a large beetle captured in a mist net. 






DMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



57 



Breeding — A male taken on 2 November in the 
Marosohy Forest had testes that were 12X8 mm 
(left) and 8X4 mm (right). A fledged juvenile 
was noted at 440 m along the trail between Isaka- 
Ivondro and Eminiminy in early January. 

Soft Part Colors — Bill: dark brown or horn 
brown with dull yellow flange; mouth lining: dull 
yellow; legs: dirty brownish yellow to golden 
brown merging to greenish yellow toes; claws: 
brownish yellow or dull greenish yellow; iris: 
brown. 

Weight— Combined (2), 183, 186 g. 

Local Name — Pokafo (Marosohy). 



Brachypteracias squamiger 
Scaly Ground-Roller 

The Scaly Ground-Roller was only noted in ar- 
eas with relatively intact closed-canopy humid 
forest between 70 and 525 m. Records from in or 
near parcel 1 of the RNI d'Andohahela include 
several individuals along the Enakara-Antseva 
trail between 325 and 425 m; one bird along the 
Isedro Trail, east of the Col d'Ambatomaniha, at 
about 525 m; and at least six pairs along 2.3 km 
of trail at our 1995 camp at 440 m. Feather re- 
mains of one individual were found in a rock shel- 
ter along the Isedro Trail at 425 m and were prob- 
ably from an individual hunted by humans. A 
specimen was taken near Eminiminy on 8 October 
by Bluntschli (smf). This species also was ob- 
served in the Manantantely Forest at 70 m. 

Brachypteracias squamiger has a call similar to 
that of B. leptosomus, but the whistle is somewhat 
higher in pitch and repeated only every 5-10 sec- 
onds. The song is given from perches 0.1-5 m 
high. No evidence of countersinging between ter- 
ritorial individuals was found in this species. 

Breeding — A male taken on 21 October at 440 
m in parcel 1 of the RNI d'Andohahela had testes 
5X4 mm (left) and 4X3 mm (right). 

Soft Part Colors — Bill: dull blackish brown 
with slightly lighter cutting edge; legs: pinkish or- 
ange; claws: dull white; iris: brown; fleshy eye 
ring: orangish pink. 

Weight— Male (1), 155 g. 

Local Name — Tatarobobaky na lakopaky (par- 
cel 1 of the RNI d'Andohahela). 

Atelornis pittoides 

Pitta-like Ground-Roller 

The Pitta-like Ground-Roller was observed at 
scattered localities in and near parcel 1 of the RNI 



d'Andohahela, all of which were in pristine or 
relatively intact humid forest. Records include in- 
dividuals on the Entseva-Anakara and Anakara- 
Ranomafana-Sud trails between 800 m and 1150 
m, one bird on the west side of the Col 
d'Ambatomaniha at about 1100 m, and at least 
four pairs along 1 km of trail just above our 1995 
camp at 1200 m. Along this trail it was sympatric 
with A. crossleyi. This site and PN de Ranoma- 
fana (M. Putnam, pers. comm.) are the only 
known localities where the two Atelornis species 
share adjacent or overlapping territories. A. pit- 
toides was not recorded anywhere else along the 
1995 altitudinal transect in the RNI d'Andohahela 
except in transitional forest north of Trafonaomby 
at 1500-1700 m, where it also was very common 
and where A. crossleyi was apparently absent. 

Breeding — Two males taken on 9 November 
had testes (left and right, respectively) that were 
6X4 mm and 5X3 mm, and 7X5 mm and 6 
X 3 mm. These two specimens did not have brood 
patches. 

Soft Part Colors — Bill: black; legs and 
claws: dull pinkish gray; iris: brown. 

Weight— Male (2), 85.5, 89.5 g. 

Local Name — Fantsasatry (Marosohy). 



Atelornis crossleyi 

Rufous-headed Ground-Roller 

The only forest in which we found the Rufous- 
headed Ground-Roller was parcel 1 of the RNI 
d'Andohahela, within an elevational range of 
800-1800 m. During the 1995 inventory of the 
reserve, this species was common at 1200 m and 
1500 m. The highest densities were at 1500 m, 
with at least eight pairs along 2 km of trail. Single 
pairs or individuals were also found at 810 and 
1800 m. Up to three singing males were audible 
from one point-count site at 1500 m. 



Leptosomatidae 

Leptosomus discolor [discolor] 
Cuckoo Roller 

The Cuckoo Roller occurs from the Marovony 
Forest south through the Anosyenne and Vohi- 
mena mountains to Petriky and west across the 
spiny forest to the Mandrare River. This species 
was occasionally heard in Tolagnaro. It was found 
at elevations between near sea level and 1800 m. 



58 



FIELDIANA: ZOOLOGY 



The Cuckoo Roller occurs in a variety of habitats 
from pristine to slightly disturbed humid, littoral, 
and spiny and gallery forest to heavily degraded 
secondary forest, transitional habitat between dry 
and humid forest, open areas at the forest edge, 
and occasionally agricultural areas several kilo- 
meters from forest. This species also moves be- 
tween habitats. For example, on 28 December 
1992 a Cuckoo Roller was observed flying and 
calling over the humid/transitional forest on the 
west side of parcel 1 of the RNI d'Andohahela, 
below the Col d'Ambatomaniha, and this individ- 
ual continued flying down the slope into spiny 
forest a few kilometers away. 

Although relatively common in humid forests, 
this species was rare in littoral forest, where our 
only record is from the Petriky Forest. Specimens 
from potential coastal areas include one taken 7 
km north of Tolagnaro (mnhn). The distinctive 
call of this species was often our means of rec- 
ognizing its presence in a given area, and thus 
lack of records from some forest sites may be 
more a function of the time of year birds vocalize 
than of their actual distribution. 

Occasionally Leptosomus will perform the ae- 
rial display in groups. On 7 June about 10 indi- 
viduals were seen and heard flying high over the 
RP de Berenty, and on 18 November in parcel 1 
of the RNI d'Andohahela, four males (including 
one subadult) spent 25 minutes continuously call- 
ing and flying over a tree in which a female was 
perched. An interesting aspect of this display is 
that the female appears to call using the same 
phrase as the males. 

Diet — The general hunting technique involves 
perching motionless, watching, and then sally- 
gleaning insects, caterpillars, and small verte- 
brates off vegetation substrate. An individual 
taken 7 km north of Tolagnaro (mnhn) on 9 March 
had debris of insects, including cicadas and grass- 
hoppers, in its stomach. 

Breeding — A nest of Leptosomus placed in a 
Neotina isoneura tree was found in the RP de 
Berenty. The specimen noted above from north of 
Tolagnaro had no enlarged ovarian follicles. 

Local Names — Vorondreo and treo-treo (Ber- 
enty). 

Upupidae 

Upupa epops marginatus 
Hoopoe 

In southeastern Madagascar the Hoopoe was a 
locally common inhabitant of open or disturbed 



lowland areas; in other areas of the island this 
species is known to occur up to 1500 m (Lan- 
grand, 1990). Our northernmost record along the 
seacoast is from the Manafiafy region. The vast 
majority of our records are either from open areas 
along the coastal plain and adjacent to littoral for- 
est, such as Manafiafy, Mandena, Itapera, and Pe- 
triky, or from spiny forest and gallery forest. We 
found no evidence that this species inhabits humid 
forest. In dry areas it appears to prefer slightly 
degraded and more open forests such as Bealoka 
and Berenty. There are three specimens taken at 
Amboasary-Sud in late October and mid-Novem- 
ber (amnh, smf). The form occurring in the area 
is U. e. marginatus. On the basis of plumage and 
song, this subspecies is markedly different from 
other forms of U. epops and probably warrants 
recognition as a distinct species. 

Diet — Chicks in a nest at Bealoka were fed 
crickets and beetles by the adults. 

Breeding — Active nests have been found at 
Bealoka in Tamarindus trees on 9 October and 14 
November. One nest had six eggs. Two specimens 
taken on 18 September at Mandena included a 
male with testes 8X4 mm and a female with a 
10 X 8 m ovary and ovarian follicles up to 2 mm 
in diameter; another pair taken on 8 October in 
the Ankapoky Forest had small gonads. 

Weight— Female (2), 57, 72 g; male (2), 69, 
89 g; combined (5), 72.4 ±11.5 (57-89) g. 

Soft Part Colors — Bill: black or dark gray at 
base merging to black tip; legs: light to dark gray- 
ish brown or dull black; claws: black; iris: dark 
brown. 

Local Name — Tsikodara (Bealoka, Berenty, 
Manafiafy, Marosohy). 



Passeriformes 

Eurylamidae 

Philepitta castanea 
Velvet Asity 

The Velvet Asity was recorded at a variety of 
sites, all of which are humid forests. This species 
was relatively common in the Marosohy Forest 
between 350 and 1000 m and in parcel 1 of the 
RNI d'Andohahela between 440 and 1900 m. Re- 
cords from other areas of parcel 1 include along 
the Tanatana Trail at about 600 m and on the east 
side of the Col d'Ambatomaniha at about 1 100 m. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



59 



Our only other observations of it from southeast- 
ern Madagascar are a pair in the Marosalohy For- 
est at about 1 1 25 m and an adult male netted at 
the edge of the Marovony Forest in a low-lying 
area with secondary scrub and Ravenala at about 
50 m. Appert (1985) reported it from the forest 
of Bemangidy. One specimen was collected in 
1756 near Fort-Dauphin (Stresemann, 1952). 

Diet — The stomachs of two individuals taken 
30 km NNW Fort-Dauphin (mnhn) contained 
seeds, vegetable debris, and fruits. In the RNI 
d'Andohahela (parcel 1) this species was noted 
feeding on the red berries of Oncostemon. 

Breeding — Several specimens taken in the Ma- 
rosohy Forest in late November and early Decem- 
ber were in or approaching breeding condition 
based on testes size. This group included males in 
breeding plumage with well-developed wattles 
and brightly colored soft parts, females about to 
lay eggs, and males in "female" plumage. Several 
males in breeding plumage and none of the males 
in female plumage had defined brood patches. Lit- 
tle is known about the breeding system of this 
species, although Langrand (1990) mentioned that 
both parents feed the young. 

In the RNI d'Andohahela (parcel 1), a female- 
plumaged bird was recorded nest building at 810 
m on 2 November. The nest was made of long 
strips of moss or lichen woven into a 10-cm-di- 
ameter ball suspended from a 1 -cm-diameter 
branch about 3 m off the ground. On 1 9 Novem- 
ber at 1500 m in the same reserve, a female-plum- 
aged individual was seen gathering nest material, 
also 5-10-cm-long strips of moss or lichen. 

Several apparent leks of this species were re- 
corded in parcel 1 of the RNI d'Andohahela. On 
1 1 November at 1 200 m, three male-plumaged 
birds spent 20 minutes calling vigorously in an 
area (not otherwise distinguishable from sur- 
rounding vegetation) about 10 X 10 m. Two males 
had completely black plumage and large wattles, 
and one male had some yellow fringes on the 
mantle and belly feathers and only small wattles. 
A fourth bird in female plumage also was present 
and chased and fought with one of the black 
males. This bird may have been a female-plum- 
aged male. One of the completely black birds had 
a much larger wattle than the other; the lobes of 
the wattle almost touched over the head and ex- 
tended to within 2 mm of the tip of the bill. This 
bird fought vigorously with the other black-plum- 
aged bird, which apparently chased the large-wat- 
tled bird away. 

On 8 November at 1200 m in the same reserve, 



two birds in female plumage chased each other 
through the understory and fought briefly. One of 
these birds had a single tertial feather with a black 
outer web. This feather also had a yellow fringe. 
On 20 November at 1 500 m, two black-plumaged 
males perched on branches 2 m from each other, 
and each bird flicked both wings slightly out and 
away from the body, exposing the yellow feath- 
ering on the carpal joint. After about 5 minutes 
of wing-flicking and singing or calling, they 
fought briefly in the middle of a bush before one 
bird flew about 20 m away. Both birds had small- 
er (but normal-sized) wattles compared with those 
of the male seen on 1 2 November. 

Weight— Female (9), 36.6 ± 1.9 (33.5-39.5) 
g; male (8), 33.4 ± 4.0 (25.5-38.5) g; combined 
(51), 35.7 ± 3.9 (25.5-47.0) g. 

Soft Part Colors — Adult Male Breeding 
Plumage — Bill: black often with yellow flange at 
base of gape; legs: yellowish green or greenish 
brown merging to yellowish green feet; claws: 
gray; iris: dark brown; fleshy caruncle: brilliant 
lime green with brilliant electric blue line above 
eye and black underside. One male in breeding 
plumage had a fleshy wattle greatly reduced in 
size and colored dark olive green above eye and 
the balance was black. Birds in Female Plum- 
age — Bill: dark brown or black with yellow flange 
at base of gape; mouthparts: yellow; legs: olive 
brown, yellowish green, or greenish brown merg- 
ing to yellowish green feet; claws: dark gray; iris: 
dark brown; orbital ring: sometimes slightly de- 
veloped and yellowish green or dull lime green. 

Local Name — Asity (parcel 1 of RNI 
d'Andohahela). 



Neodrepanis coruscans 
Sunbird-Asity 

The only localities where we found this species 
were in the Marosohy Forest and the adjacent RNI 
d'Andohahela (parcel 1) between 425 and 1350 
m. The lower elevation records include a bird in 
the Marosohy Forest at 425 m and a male west 
of Eminiminy between 440 and 600 m. The low- 
est elevation at which this species was recorded 
in the RNI d'Andohahela was 810 m. At 1350 m, 
it was recorded in forest about 50 m elevational 
distance below where N. hypoxantha were seen 
regularly; N. hypoxantha was along the crest of a 
ridge, whereas N. coruscans was present in adja- 
cent contiguous forest on the slopes of the same 
ridge. Hence the altitudinal separation of these 



60 



FIELDIANA: ZOOLOGY 






two species was sharply defined. Appert (1985) 
reported a Neodrepanis, presumably coruscans, 
from the Bemangidy Forest. 

Several birds in the RNI d'Andohahela (parcel 
1) at 810 m were in immature male plumage, with 
a few blue-fringed feathers on the scapulars or 
coverts and rump, a small ( 1 mm diameter) bluish 
wattle over the eye, and dull yellow underparts. 
Some of these individuals may have been males 
in "female" plumage, a direct parallel to the sit- 
uation already described for Philepitta. Also pres- 
ent in the same area at the same time were males 
in full breeding plumage. 

Males when flying make a noticeable whistling 
or humming sound with their wings. This sound 
is not produced by female or immature male N 
coruscans, and it is not as noticeable as in male 
N hypoxantha. In many bird species such me- 
chanical noises are produced by modified (often 
highly emarginated) outer primaries. The first 
(outermost) primary in male N. hypoxantha is 
broadly emarginated, whereas no such broad 
emargination is apparent in the primaries of adult 
male N coruscans. 

Diet — In the Marosohy Forest at about 900 m 
an adult male Sunbird-Asity was observed feeding 
at the flowers of Sloanea. The stomach contents of 
one bird consisted of small insects and some liquid 
that may have been flower nectar. A female at 810 
m fed in a clump of Bakerella flowers. 

Breeding — An adult male in breeding plumage 
taken on 25 November had a well-developed ca- 
runcle and brightly colored eye ring but small tes- 
tes. All males in breeding condition lack brood 
patches. A female at 810 m on 2 November was 
collecting nest material (short strips of lichen or 
moss), and a male at 810 m on 6 November, 
which flew back and forth across a river with 
food, presumably was feeding young. 

Soft Part Colors — Bill: black except at base 
of mandible, which varies from dark green to bril- 
liant green; legs and claws: black; iris: brown; 
fleshy area around eye: deep cobalt blue (males 
in breeding condition). 

Weight— Female ( 1 ), 7.0 g; male (4), 6.5 ± 0. 1 
(6.4-6.7) g; combined (5), 6.6 ± 0.2 (6.4-7.0) g. 
Local Name — Soy (Marosohy). This is the 
same name used for Nectarinia spp.; the infor- 
mant may have confused these two genera. 

Neodrepanis hypoxantha 

Yellow-bellied Sunbird-Asity 

Our only records of the Yellow-bellied Sunbird- 
Asity are from parcel 1 of the RNI d'Andohahela 



between 1350 and 1950 m, in which zone it was 
common. This species was absent from transition- 
al forest at 1700 m north of Trafonaomby and 
along the western slope of the main massif. In late 
May one individual was observed in the RNI 
d'Andohahela between Esomony and Vohibaka at 
1 150 m (Langrand & Sinclair, 1994; OL). 

A description of juvenile birds has been pre- 
sented by Hawkins et al. (in press). Birds appar- 
ently having completed postjuvenile molt were 
noted on several occasions at 1500 m (20 and 22 
November) and 1900 m (28 and 30 November). 
These birds resembled female N. coruscans in that 
they had dull olive underparts except for a pale 
yellow area on the flanks. The wing coverts had 
a narrow pale buff fringe. The major distinction 
between N. hypoxantha and N. coruscans at this 
age is the call, which in the former species is 
much higher pitched and generally a single note. 
Diet — At 1500 m, female and immature male- 
plumaged birds both fed from the long-spurred 
flowers of Bakerella. The birds did not use the 
length of their bill to reach the nectar source but 
instead inserted their very long tongues. Other 
food plants noted (both flowers apparently visited 
for nectar) included Gaertnera and Medinilla. 

On 1 December, at 1960 m on the summit of 
Trafonaomby, a male N. hypoxantha disputed ac- 
cess to a clump of Bakerella flowers with two 
female Nectarinia souimanga. It was difficult to 
tell the outcome of the dispute, but the N. soui- 
manga were continually present at the site and 
Neodrepanis hypoxantha appeared to be only a 
casual visitor, suggesting that the sunbird was the 
dominant species. 

Breeding — A bird netted on 22 November in 
female plumage had a brood patch, and a speci- 
men collected on 1 December had only 15% of 
the skull ossified. Males in breeding plumage 
lacked brood patches. In the RNI d'Andohahela 
(parcel 1) a nest was found on 2 November at 
1850 m that contained two eggs. The nest was 
oval in shape and suspended from a lateral branch 
of a 2.5-m sapling on a steep slope. The nest was 
collected on 4 December and the eggs contained 
small embryos. Only the female was seen brood- 
ing the eggs, but when the nest was collected both 
the male and the female vigorously mobbed the 
intruder. 

On 19 November at 1500 m an immature male- 
plumaged bird (with individual blue-fringed black 
feathers on lower and upper scapulars and tertials 
and a half-sized blue wattle) displayed to a fe- 
male-plumaged bird by fluffing out the body 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



61 



feathers, drooping the bill and head forward, and 
depressing the tail. During this display the male 
called vigorously, a series of single hissing notes. 

Soft Part Colors — Adult Male in Breeding 
Plumage — Bill: at base dark cobalt blue posterior 
to nostril, light cobalt blue above (dorsally) and 
connecting nostrils, bright lime green around na- 
sal operculum, and balance of bill black; legs: 
brownish black; claws: black; iris: dark brown; 
fleshy caruncle: complex pattern of color and 
slightly variable with the lower half (below eye) 
dark cobalt blue, lateral dull dark green stripe the 
width of the eye and directly posterior to it, above 
eye light cobalt blue, sometimes bright sky blue, 
and area between eye and base of bill dark cobalt 
blue. A male molting into breeding plumage had 
a dull cobalt blue fleshy caruncle and some bril- 
liant blue and green skin around the nostrils. Fe- 
male Plumage — Bill: dull brownish black; legs: 
dark grayish brown or brownish black; foot pads: 
dull yellow; claws: black; iris: dark brown. 

Weight— Presumed female (3), 7.0 ± 1.0 (6.0- 
8.0) g; male (3), 7.5 ± 0.5 (7.0-8.0) g; combined 
(8), 7.2 ± 0.8 (6.0-8.0) g. 



Alaudidae 

Mirafra hova 

Madagascar Bush Lark 

The Madagascar Bush Lark was common in de- 
graded open areas from the Marovony Forest south 
through the Anosyenne and Vohimena mountains 
to Petriky and west to the Mandrare River from 
near sea level to over 1700 m. This species was 
abundant along the eastern coastal plain in open 
grasslands, agricultural fields, heathlands, and de- 
graded areas near the littoral forests of Manafiafy, 
Itapera, Mandena, and Petriky. Further inland, in 
regions with lateritic soils it was less common, al- 
though it was locally common in some upland ar- 
eas, such as agricultural fields in the Manampanihy 
River valley between the Col de Managotry and 
Ranomafana-Sud. During the 1995 inventory of 
the RNI d'Andohahela (parcel 1), this species was 
not found in humid forest and was only present in 
open grassland adjacent to the transitional forest 
between 1500 and 1700 m. In the spiny forest re- 
gion it occurs in a variety of habitats from degrad- 
ed and slightly open areas in spiny forest to river 
plains, degraded areas, and sisal plantations. 

Diet — Stomach contents of collected birds in- 
cluded beetles (Curculionidae, Elateridae, Scara- 



baeidae, Tenebrionidae), ants (Formicidae), and 
flies (Asilidae) (Goodman & Parrillo, in press). 

Breeding — A male and female taken near 
Manafiafy in the first half of October had testes 6 
X 4 mm and a slightly enlarged ovary, with thick- 
ened oviduct and ovarian follicles up to 3 mm in 
diameter, respectively; only the female had a 
brood patch. Also, during the same period another 
collected bird had a partially ossified skull and 
small gonads. A male taken at Ankapoky on 12 
October had testes 3X2 mm and no brood patch. 

Weight— Combined (5), 19.6 ± 2.2 (17.5- 
23.0) g. 

Soft Part Colors — Bill, legs, and claws: light 
pink to horn gray; iris: brown. 

Local Names — Borisy (Marosohy), bria (Man- 
ombo), jorioke (Berenty). 

Hirundinidae 

Riparia paludicola [cowani] 
Brown-throated Sand Martin 

The only known record of the Brown-throated 
Sand Martin in southeastern Madagascar is five 
individuals flying over a marsh surrounded by 
transitional forest at 1 500 m, north of Trafonaom- 
by. In other parts of Madagascar, it is rare below 
elevations of 500 m (Langrand, 1990). The field 
work conducted in the southeast was generally in 
lowland areas, which might account for the lack 
of sightings elsewhere in this region. 



Phedina borbonica madagascariensis 
Mascarene Swallow 

This species was recorded sporadically between 
sea level and 1950 m elevation. It often is a hu- 
man commensal, nesting in abandoned buildings 
and various types of structures. It was common 
along the coastal plain between Manantenina and 
Tolagnaro, where small groups were observed for- 
aging over villages and agricultural areas. We re- 
corded this species on numerous occasions in the 
interior of large intact humid forests, e.g., the 
Marosohy Forest and parcel 1 of the RNI 
d'Andohahela. In the spiny forest region it was 
distinctly less common, except for areas along riv- 
ers and near villages or towns, such as the RP de 
Berenty, Ambovombe, and Amboasary-Sud. We 
found no evidence of this species breeding along 
natural rock faces, in caves, or in tree cavities; in 
southeastern Madagascar it clearly favors human- 
made structures for nesting sites. 



62 



FIELDIANA: ZOOLOGY 



This species was often observed in mixed-spe- 
cies flocks with other aerial foraging insectivores, 
such as Zoonavena, Apus barbatus, A. melba, 
Cypsiurus, and Hirundo rustica. 

Diet — A specimen taken in the Tolagnaro area 
had beetle (Elateridae, Scarabaeidae), Homoptera, 
and ant (Formicidae) remains in its stomach 
(Goodman & Parrillo, in press). 

Breeding — Different active nests were found in 
Tolagnaro: on 14 September with four eggs, on 
17 September with two eggs, on 6 November with 
three nestlings about 10 days old, and on 15 Oc- 
tober with four eggs. Collected females in breed- 
ing condition had brood patches, whereas males 
lacked brood patches. All breeding sites found to 
date in the region are in human-made structures. 

Weight— Combined (4), 21.3 ± 1.0 (20.0- 
22.5) g. 

Soft Part Colors — Bill: black; legs: dark 
brown; claws: black; iris: brown. 

Local Names — Fililiotra (Marosohy), tsilotsi- 
lon'aka (Manafiafy); both are generic for swal- 
lows and swifts. 



Hirundo rustica subsp. 
Barn Swallow 

This species is a relatively uncommon migrant 
to Madagascar during the boreal winter (Lan- 
grand, 1990). Our only record of it in southeastern 
Madagascar is one observed at Tolagnaro on 12 
November in a flock of Phedina borbonica. 



Motacillidae 

Motacilla flaviventris 
Madagascar Wagtail 

The Madagascar Wagtail was infrequently ob- 
served from the Marovony Forest south through 
the Anosyenne and Vohimena mountains and 
along the coastal plain north to the Manantantely 
Forest. Its elevational range in the region is from 
sea level to 1950 m. Most records are from wet 
open areas, such as pastureland, rice paddies, and 
the margins of rivers and marshes. In humid forest 
(e.g., parcel 1 of the RNI d'Andohahela) it is al- 
most always along streams. We found this species 
near or adjacent to the forest parcels of Marovony, 
Analalava, Marosohy, Manafiafy, Itapera, Man- 
dena, Bezavona, and Manantantely. We observed 
it on a few occasions in eucalyptus plantations 



bordered by small streams. West of the Anosyen- 
ne Mountains, in the spiny forest region, it was 
distinctly rare and generally confined to the mar- 
gins of permanent rivers such as the Mandrare or, 
in the RP de Berenty, open areas adjacent to the 
tourist bungalows. 

Salvan (1970) reported that this species may be 
a seasonal migrant in the southern portion of the 
island, being absent during the austral winter. 
During the period of the QIT field studies, Sep- 
tember through December (austral summer), we 
saw no change in the abundance of this species. 

Diet— Individuals collected in 1989 and 1990 
had the following food remains in their stomachs: 
beetles (Curculionidae, Elateridae), Homoptera, 
and Hymenoptera (Goodman & Parrillo, in press). 
This species was observed along the coastal beach 
feeding on insects in tidal flotsam. 

Breeding — In mid-October an adult Madagas- 
car Wagtail was observed carrying food, presum- 
ably to a nest. Specimens taken at Manafiafy in 
late October were an immature with small testes 
and a bursa 6X2 mm and an adult male with 
testes 6X4 mm and no brood patch. A male 
taken at the edge of the Marovony Forest on 30 
October had testes 4x3 mm and a cloacal pro- 
tuberance, and a female taken at the same time 
had a slightly enlarged ovary and a brood patch. 
On 31 October we observed an adult bringing 
food to a nest placed in a crevice of a large boul- 
der resting in the middle of a stream along the 
Enakara-Antseva trail (325 m elevation). The 
stream formed the boundary between the Maro- 
sohy Forest and a rice paddy. A similarly placed 
nest was found along the Andranohela River at 
about 400 m. 

Soft Part Colors — Bill, legs, and claws: 
black or brownish black; iris: dark brown. 

Weight— Combined (10), 22.3 ± 0.9 (21-24) g. 

Local Name — Triotho (Berenty, Manafiafy, 
Manombo, Marosohy). 

Campephagidae 

Coracina cinerea cinerea and Coracina 
cinerea pallida 

Ashy Cuckoo Shrike 

The Ashy Cuckoo Shrike was regularly noted 
in primary and secondary humid forest from the 
Marovony Forest south through the Anosyenne 
and Vohimena mountains to Manantantely, from 
20 to 1950 m. During the 1995 inventory of par- 
cel 1 of the RNI d'Andohahela this species ap- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



63 



peared equally common at all altitudinal zones 
sampled between 440 and 1950 m. Appert (1985) 
reported this species from near Bemangidy. It also 
has been collected 30 km NNW Fort-Dauphin 
(mnhn) and near Eminiminy (amnh). The only 
record we have from littoral forest is a pair at 
Manafiafy on 10 October (fmnh), although this 
species was observed in or near Mandena (GR). 
A specimen was collected in 1756 near Fort-Dau- 
phin (Stresemann, 1952). All material examined 
from the Anosyenne Mountains and east to the 
coast is referable to C. c. cinerea. The Ashy 
Cuckoo Shrike also occurs in riparian habitat, 
such as the RP de Berenty, Bealoka, and Hazo- 
fotsy and along the Mananara River, where the 
subspecies presumably is C. c. pallida. 

In early October 1995 feathers of this species 
were found along the Tanatana Trail between Is- 
aka-Ivondro and Eminiminy. These remains were 
presumably those of a hunted individual killed 
with a slingshot. 

Diet — Stomach contents of birds we collected 
included Diplopoda, spiders (Araneae), beetles 
(Curculionidae, Elateridae, Scarabaeidae, Teneb- 
rionidae), Lepidoptera (larvae), Isoptera, Homop- 
tera, and Orthoptera (Tettigoniidae) (Goodman & 
Parrillo, in press). A large millipede was found in 
the stomach of one bird (mnhn). The stomach 
contents of a bird collected in the Analalava For- 
est from a mixed-species foraging group consisted 
of fruits and parts of at least one orthopteran. Ob- 
servations of foraging individuals include a male 
feeding on a 30-mm caterpillar, an unsexed bird 
foraging on small fruits in a tree in the Manan- 
tantely Forest, and an adult dispatching a praying 
mantid in the Bealoka Forest. Feeding behavior 
often involves probing trees with cracked or con- 
voluted bark, such as Euphorbia tirucalli, Fer- 
nandoa madagascariensis, and Tamarindus. 

Breeding — A pair taken at Manafiafy on 10 
October consisted of an adult male with gray tes- 
tes 7 X 4 mm and an adult female with an ovary 
7X6 mm and with no greatly enlarged ovarian 
follicles. A female obtained on 28 September in 
the Manantantely Forest had an enlarged ovary, a 
partially shelled egg in the oviduct, and two cor- 
pora lutea. 

On 17 November at 1200 m in parcel 1 of the 
RNI d'Andohahela, a male and a female were dis- 
playing to each other. The display consisted of a 
rapid series of short wing flicks, either a single 
wing or both wings together, while the birds gave 
a repeated quiet whistle. In the same reserve, a 
male and a female shared duties at a nest at 1 650 



Table 5. Number of Phyllastrephus madagascar- 
iensis, P. zosterops, and P. cinereiceps netted at humid 
forest sites. 





No. of 










accrued 


P. 




P. 


Site, 


net- 


madagas- 


p. 


ciner- 


Elevation 


days 


cariensis 


zosterops 


eiceps 


Analalava 










40 m 


31 


11 


2 





Marosohy 










425 m 


50 


7 


3 





725 m 


63 


4 


12 





Marovony 










50 m 


110 


16 








Manantantely 










100 m 


32 


7 








Andohahela 










440 m 


50 


8 


4 





810 m 


50 


1 


3 





1200 m 


50 


3 


7 


1 


1500 m 


50 








4 


1875 m 


50 












m in late November. The nest was a shallow cup 
made of flaky lichens, held together with spider 
web, with a lining made of moss. It was situated 
on a 1 0-cm-diameter horizontal branch about 7 m 
above the ground. 

Weight— Combined (6), 47.0 ± 2.9 (42.5- 
51.0) g. 

Soft Part Colors — Bill, legs, and claws: 
black; iris: dark brown. 

Local Names — Bokazava (Manantantely), bok- 
azavo (Manombo), mavoloha (Berenty), tsirikiti- 
tio (Berenty). 



Pycnonotidae 

Phyllastrephus madagascariensis 
madagascariensis 
Long-billed Greenbul 

This species was found in humid forests from 
the Marovony Forest south through the Anosyen- 
ne and Vohimena ranges to Manantantely between 
50 and 1200 m. During the 1995 inventory of 
parcel 1 of the RNI d'Andohahela, this species 
was found between 440 and 1200 m, although it 
was markedly less common at 1200 m than at 
lower elevations (Table 5). At 1200 m it was 
found only in valleys where the vegetation resem- 



64 



FIELDIANA: ZOOLOG" 



bled that of lower elevations. Appert (1985) re- 
ported it from the Bemangidy Forest. It was not 
recorded in heavily disturbed forested areas or in 
littoral forest. All material examined from the re- 
gion is referable to the nominate subspecies. 
There is an unsubstantiated report of this species 
in the Ankoba Forest, near the RP de Berenty 
(Rakotondranony, 1977); this forest is primarily a 
monoculture of Pithecellobium dulce, and without 
further details this record cannot be accepted. 

Diet — Stomachs examined contained spiders 
(Araneae), Diplopoda, Blattodea, Coleoptera 
(Chrysomelidae, Cleridae, Curculionidae, Elater- 
idae, Tenebrionidae), Homoptera (Cicadellidae, 
Cicadidae, Fulgoroidea), Hymenoptera (Formici- 
dae), Orthoptera, and Reptilia (Gekkonidae) 
(Goodman & Parrillo, in press). 

Breeding — Specimens taken in the Analalava 
Forest between 5 and 10 November were mostly 
adults in reproductive condition. Adult males 
lacked or had indistinct brood patches, whereas 
adult females often had well-defined or vascular- 
ized brood patches. The only nonadult taken dur- 
ing this period was a bird with a totally unossified 
skull and a 3-mm bursa. Seven birds captured in 
the Manantantely Forest between 30 September 
and 2 October, 1 1 birds handled in the Marosohy 
Forest between 29 November and 1 1 December, 
and 15 birds netted in the Marovony Forest be- 
tween 26 October and 3 November were all 
adults. On 27 December in the RNI d'Andohahela 
(parcel 1 ) at about 900 m we saw an adult feeding 
two fledglings, and at about 950 m we saw two 
immatures unaccompanied by adults. 

Weight— Female (12), 25.4 ± 2.7 (22.0-31.5) 
g; male (15), 32.0 ± 4.3 (22-39) g; combined 
(56), 30.8 ± 4.4 (22-39) g. 

Soft Part Colors — Bill: maxilla dark gray or 
dark brown and in subadults dusky, and mandible 
(highly variable) horn gray at base merging to 
dark brown tip, dull pinkish brown or pale gray 
brown with dusky tip and in subadults medium 
brown and medially yellowish brown; mouth lin- 
ing: yellow; legs: grayish brown or brown and 
sometimes with orange tinge; foot pads: ochre 
yellow; claws: gray; iris: medium brown to dark 
brown and in subadults gray brown. 

Local Names — Parataky (Marosohy). pretaka 
(Marovony), pretaky (Manombo). 



Phyllastrephus zosterops zosterops 
Spectacled Greenbul 

The Spectacled Greenbul was relatively com- 
mon in the closed-canopy humid forests of Ma- 



rosohy, Analalava, and Manantantely. This spe- 
cies also was found in various areas of the RNI 
d'Andohahela (parcel 1) between 440 and 1200 
m elevation; it was much less common at 1200 m 
than at lower altitudes (Table 5). It also was re- 
corded in transitional forest north of Trafonaomby 
between 1500 and 1700 m. At all of these local- 
ities except the last it was sympatric with P. mad- 
agascariensis. At a few sites with apparently sim- 
ilar forest types (e.g., Marovony and Manantan- 
tely forests), however, madagascariensis was rel- 
atively common and zosterops was absent. P. 
zosterops was not recorded in littoral forest. All 
specimens examined from the region are referable 
to the nominate form. 

Diet — The stomach contents of collected birds 
contained spiders (Araneae), beetles (Curculioni- 
dae, Elateridae, Hydrophilidae, Scarabaeidae, 
Staphylinidae, Tenebrionidae), Dermaptera, Isop- 
tera, flies (Nematocera), Homoptera, ants (For- 
micidae), and Orthoptera (Tetrigidae) (Goodman 
& Parrillo, in press). 

Breeding — Most of the individuals collected in 
the Analalava and Marosohy forests between ear- 
ly November and mid-December were adults in 
reproductive condition, including males with tes- 
tes up to 1 1 X 7 mm and females with enlarged 
ovaries and ovarian follicles up to 9 mm in di- 
ameter. Males apparently lack brood patches. 

Many nests and fledged juveniles of this spe- 
cies were found in parcel 1 of the RNI 
d'Andohahela between 18 October and 6 Novem- 
ber. All nests were deep cups, suspended between 
the branches of understory shrubs or, in one case, 
2 m above the ground in a Diospyros sapling. 
Nests with two fledglings, in each case estimated 
to be 2 or 3 days old, were found at 440 m (one 
nest) and 810 m (two nests). At 440 m, seven 
groups of adults feeding juveniles were seen: four 
with one juvenile, one with two, and one with 
three. In two cases, three adults were feeding the 
juveniles. The extra individual in each case may 
have been a "helper." Apparent helpers also have 
been noted in P. cinereiceps (see below; also 
Hawkins et al., in press). On 29 November a pair 
of adults was feeding two newly fledged juveniles 
in transitional forest at 1700 m, north of Trafon- 
aomby. 

Weight— Female (5), 17.7 ± 2.9 (15.5-22.5) 
g; male (6), 19.0 ± 1.0 (18.5-20.5) g; combined 
(35), 18.3 ± 1.8 (13.5-22.5) g. 

Soft Part Colors — Bill: maxilla brown with 
yellow cutting edge or completely black, and 
mandible yellow or orangish yellow sometimes 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



65 



with brown tip; mouth lining: bright yellow; legs: 
grayish pink, grayish brown, or grayish yellow; 
foot pads: yellow; claws: pinkish gray or gray; 
iris: brown. 



Phyllastrephus cinereiceps 
Gray-crowned Greenbul 

The Gray-crowned Greenbul has been recorded 
in southeastern Madagascar only in the humid for- 
est of the RNI d'Andohahela between 810 and 
1950 m. At 810 m this species was only found in 
a small area with elements of upper montane for- 
est. At higher elevations it was more common, 
although less so above 1850 m in low mossy for- 
est near the summit of Trafonaomby (Table 5). It 
appeared to be absent from the transitional forest 
between 1500 and 1700 m north of Trafonaomby. 

Breeding — On 13 November at 1200 m, three 
adults were feeding two recently fledged juve- 
niles. The fledglings were dull gray-green above, 
the same color on the crown, and dull buffy un- 
derneath with a slightly paler throat. At 1650 m 
on 21 November, two adults were seen with two 
juveniles. These fledglings were older than those 
seen at 1200 m and had uniform gray-green up- 
perparts with a slight gray tinge to the crown 
feathers, pale orange gapes, and slightly yellow- 
tinged underparts and were paler, almost whitish, 
on the throat. On 4 December at 1900 m, three 
adults were accompanying two juveniles. These 
juveniles appeared to be molting into adult plum- 
age, with grayish brown caps contrasting slightly 
with olive brown backs and blotchy whitish 
throats contrasting with yellow breast and belly. 

Weight— Male (1), 18.5 g; combined (4), 17.1 
± 1.1 (16.0-18.5) g. 

Soft Part Colors — Bill: maxilla black with 
gray cutting edge, and mandible dark pinkish gray 
at base merging to black tip; legs: pinkish gray; 
iris: brown. 



Hypsipetes madagascariensis madagascar- 
iensis 

Madagascar Bulbul 

The Madagascar Bulbul was one of the most 
common and widespread birds in the humid and 
littoral forests of southeastern Madagascar from 
sea level to 1500 m. This species was found at 
every site we visited from the Marovony Forest 
south through the Anosyenne and Vohimena 



mountains and along the coastal plain to the Pe- 
triky Forest and west to the Mandrare River. It 
occurs in open habitats, particularly in heavily 
disturbed areas such as grasslands and agricultural 
fields with scattered trees, and at the forest edge, 
as well as in the heart of intact forests. West of 
the Anosyenne Mountains, in spiny forest, it was 
not as abundant, although even in this region it 
was common in relatively intact and disturbed 
spiny forest and gallery forest and along river 
margins. 

Diet — Food remains in the stomachs of col- 
lected individuals consisted of fruits (small Fi- 
cus), small seeds, spiders (Araneae), beetles (Cu- 
culionidae, Elateridae, Scarabaeidae: Hopliini), 
and Hemiptera (Reduviidae) (Goodman & Parril- 
lo, in press). They have been noted dispatching 
and feeding on large spiders such as Nephila. 
Madagascar Bulbuls have been observed feeding 
on fruits of Capparis spp., Rinoria greveana, 
Neotina isoneura, Cordia ronnii, and Celtis phil- 
lipensis and on the flowers of Crateva excelsa. 
This bulbul is able to exploit a variety of orna- 
mental plants in remote and isolated villages away 
from the forest such as Vohibaka (800 m) on the 
northern limit of RNI d'Andohahela (parcel 1), 
where it feeds extensively on the berries of intro- 
duced Melia azedarach. 

Breeding — All Madagascar Bulbuls collected 
during the QIT project were adult, and the vast 
majority were in or approaching breeding condi- 
tion. All males lacked brood patches. A nest with 
three eggs was found on 9 November at the edge 
of the Analalava Forest. The nest was placed in a 
tree about 3.5 m above the ground and secured to 
four points of two adjacent branches with spider 
web. On 15 October in the Manafiafy Forest an 
adult was observed feeding two fledglings. On 28 
November and 2 December, at different localities 
in the Marosohy Forest, adults were observed 
bringing food to nests or feeding fledglings. In the 
latter case, one adult of the pair had a large beetle 
in its beak and the other had a large moth, the 
wings of which it tore off. 

Weight— Female (7), 43.6 ± 5.8 (33-52) g; 
male (14), 44.3 ± 3.4 (39-52) g; combined (38), 
44.9 ± 4.0 (33-52) g. 

Soft Part Colors — Bill: maxilla orange often 
with dusky tip and occasionally with dusky nasal 
operculum, and mandible orange; legs: dull or- 
angish brown or dull yellowish orange; claws: 
grayish brown or black; iris: reddish brown. 

Local Names — Horova or horovana (Man- 



66 



FIELDIANA: ZOOLOG^ 



ombo, Marosohy), tsikonina (Berenty), tsikorova 
(Manantantely). 



Turdidae 

Copsychus albospecularis pica and 
Copsychus albospecularis inexpectatus 
Madagascar Magpie Robin 

The Madagascar Magpie Robin was found from 
the Marovony Forest south through the Anosyen- 
ne and Vohimena mountains to Petriky and west 
to the Mandrare River. This species was common 
in a variety of habitats, including closed-canopy 
humid forest, littoral forest, secondary natural for- 
est, degraded areas along the forest edge, gardens 
in Tolagnaro, eucalyptus plantations, intact and 
degraded spiny forest, and gallery forest between 
near sea level and 1200 m. During the 1995 in- 
ventory of parcel 1 of the RNI d'Andohahela, it 
was found only in the 440 m, 800 m, and 1200 
m elevation zones, where it was rare as compared 
with its distribution in similar habitats elsewhere 
on the island. Although this species was generally 
common in littoral forest, we did not find it at 
Mandena, and it was rare at Petriky. Copsychus 
also was not found at the Station Forestiere de 
Mandena during a survey in the 1960s or 1970s 
(GR). 

The distributions of Copsychus a. inexpectatus 
and C. a. pica were sharply defined in this region. 
Specimens (amnh, fmnh) obtained west of the An- 
osyenne Mountains, in spiny forest, are all refer- 
able to pica, as are specimens from the littoral 
forest of Petriky. Those to the east, in humid and 
littoral forest areas, are inexpectatus, including 
birds taken in the forests of Marovony, Analalava, 
Manafiafy, Bezavona, and Manantantely (fmnh) 
and 30 km NNW Fort-Dauphin (mnhn). The dis- 
tance between the populations of C. a. pica at 
Petriky and C. a. inexpectatus at Manantantely is 
less than 8 km (by air). 

Diet — The stomachs of specimens collected 30 
km NNW Fort-Dauphin (mnhn) contained insects 
and spiders. The stomach contents of our collect- 
ed material included Isopoda, Blattodea, spiders 
(Araneae), Orthoptera, Hemiptera (cf. Aradidae), 
adult beetles (Acanthoceridae, Carabidae, Cucu- 
jidae, Curculionidae, Elateridae, Scarabaeidae, 
Staphylinidae, Tenebrionidae), beetle larvae, ants 
(Formicidae), plant fibers, and small amphibians 
and geckos (Goodman & Parrillo, in press). In the 
Ankapoky Forest a mixed-species group was ob- 



served in a Dicoma; a male magpie robin and two 
Pioceus sakalava were eating blossoms, and Nec- 
tarinia notata and N. souimanga were feeding on 
the contents of flowers. C albospecularis also has 
been observed feeding on fruits and berries. 

Breeding — Most adults collected during the 
1989-1990 field seasons between late September 
and late December had enlarged gonads. A female 
obtained in the Analalava Forest on 5 November 
had an unshelled egg in oviduct, two corpora 
lutea, and an ovarian follicle 8X7 mm. Fledg- 
lings and subadults were noted in the RNI 
d'Andohahela (parcel 1) at 1200 m on 16 Novem- 
ber being fed by adult males, in the Bezavona 
Forest on 29 December, in the Marosohy Forest 
on 3 and 12 December, and in the Manafiafy For- 
est on 1 9 and 20 December. 

The 12 December fledgling had been captured 
by an Accipiter madagascariensis; the Tandroy 
people around Bealoka noted this raptor feeds ex- 
tensively on Copsychus. No adult male in breed- 
ing condition was found with a brood patch. 

Weight— C a. inexpectatus female (22), 23.7 
± 2.8 (18.5-30.5) g; male (27), 24.9 ± 1.7 (20.0- 
28.5) g. C. a. pica female (1), 18.5 g; male (1), 
19.5 g. C a. inexpectatus and pica combined (59), 
24.2 ± 2.7 (16.5-30.5) g. 

Soft Part Colors— Bill: black; mouth lining: 
dull orangish yellow (fledgling); legs: olive 
brown, light gray, or black; claws: light gray or 
black; iris: brown. C. a. pica tends to have much 
darker legs than does C a. inexpectatus. 

Local Names — Fitatra (Manafiafy), fitatr'ala 
(Manafiafy), fitatry (Marosohy), peeda (Berenty), 
tsilaka (Manombo). 



Saxicola torquata sibilla 
Stonechat 

The Stonechat was found in southeastern Mad- 
agascar from the Marovony Forest south through 
the Anosyenne and Vohimena mountains, to Pe- 
triky and west through the spiny forest to the 
Mandrare River between near sea level and 1900 
m. This species was distinctly rarer in drier por- 
tions of its distribution. Throughout this range it 
is found in open areas (often degraded), such as 
grassland and heathland, or at the edge of lakes 
and lagoons, often perched on bushes or small 
rocky outcrops. We never observed it in any type 
of natural humid forest. In the transitional forest 
north of Trafonaomby between 1500 and 1800 m, 
at least seven different pairs were located in hab- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



67 



itats ranging from edges of dense forest to open 
burnt grassland adjacent to a marsh. This species 
clearly has benefited from human-induced habitat 
modifications. We have few records of this spe- 
cies in spiny forest; one bird was observed near 
Bealoka along the river fioodplain. Specimens ex- 
amined from the area are referable to S. t. sibilla. 

Diet — The stomachs of collected individuals 
contained spiders (Araneae), Blattodea, Coleop- 
tera (Scarabaeidae, Tenebrionidae), Hymenoptera 
(Formicidae), and Orthoptera (Gryllacrididae) 
(Goodman & Parrillo, in press). 

Breeding — On 13 October at Manafiafy a pair 
of adults was collected with two fledglings. North 
of Trafonaomby, between 1 500 and 1 800 m, many 
pairs of Stonechats had fledged juveniles in late 
November and early December. 

Weight — Female (1), 16.5 g; male (1), 15.0 g. 

Soft Part Colors — Bill: black in males, dark 
brown in females, and gray in fledglings; legs: 
black or dark brown; claws: black; iris: brown. 

Local Name — Fitadroanga (Manafiafy). 



Pseudocossyphus sharpei sharpei 
Forest Rock-Thrush 

The Forest Rock-Thrush was recorded in the 
Marosohy and Marosalohy forests and parcel 1 of 
the RNI d'Andohahela, where it was observed in 
intact closed-canopy humid forest between 1100 
and 1875 m. This species also was observed be- 
tween 1500 and 1800 m in the transitional forest 
north of Trafonaomby. At this locality they fre- 
quently foraged at the edge of the forest, up to 50 
m from closed forest. Langrand (1990) reported 
this species from areas as far south as Tolagnaro 
at elevations above 810 m. The form occurring in 
the area is nominate sharpei. 

Breeding — Adult males collected in early No- 
vember had enlarged testes (maximum size: left, 
9X5 mm; right, 7X4 mm) and did not possess 
brood patches. On 1 November at 1235 m in the 
Marosalohy Forest we saw an adult female car- 
rying nesting material. Within parcel 1 of the RNI 
d'Andohahela at 1200 m on 19 November, a male 
was seen carrying food, presumably for juveniles, 
and on 2 1 November a male and female were seen 
in the company of two juveniles. At 1500-1800 
m in transitional forest north of Trafonaomby be- 
tween 27 November and 3 December three dif- 
ferent pairs were located with two, one, and one 
juveniles, and a fledgling was netted at 1875 m 
on 29 November, below the summit of Trafon- 



68 



aomby. The juveniles were in subadult plumage 
with short tails and wings, and all back and head 
feathers had paler tips. The breast feathers had 
wider pale tips. None of the wing feathers were 
tipped paler except the innermost greater coverts 
and very fine edges to the remiges. 

Soft Part Colors — Bill: black; mouth lining: 
yellow; legs: pinkish gray; feet: dull yellow; 
claws: black; iris: brown. In the fledgling the max- 
illa was black around nasal operculum merging to 
gray tip and dull yellow cutting edge and the man- 
dible was dull yellow. 

Weight— Male (6), 24.0 ± 1.4 (22.5-25.5) g; 
fledglings (2), 24.5, 24.5 g. 



Pseudocossyphus imerinus 
Littoral Rock-Thrush 

We only have a single record of this species in 
southeastern Madagascar. We saw two individuals 
on 24 December in an area of coastal dunes and 
sparse vegetation at Lac Anony, the eastern limit 
of this species' distribution (Langrand, 1990). 
Farkas's (1974) prediction that this species' range 
might extend to Tolagnaro has not been verified. 
However, what is apparently appropriate coastal 
xerophytic habitat for the Littoral Rock-Thrush 
exists along the sea south-southwest of Ranopiso, 
and this species may occur in that area. 



Sylviidae 

Acrocephalus newtoni 

Madagascar Swamp- Warbler 

Although this species is widespread across 
Madagascar (Langrand, 1990), we have few re- 
cords of the Madagascar Swamp-Warbler from 
southeastern Madagascar, despite permanent reed 
beds in the region that appear to have suitable 
habitat for this species. It has been observed near 
Saihady, 5 km north of Tolagnaro, at several sites 
along the Mandrare and Mananara rivers, along 
the Ranopiso River, and in the reed beds along 
the Tarantsy River near Bevilany. 



Cryptosylvicola randrianasoloi 
Cryptic Warbler 



The recently described (Goodman et al., 1996) 
Cryptic Warbler is known in southeastern Mada- 



FIELDIANA: ZOOLOGY 






gascar only from the RNI d'Andohahela (parcel 
1), where it was recorded between 810 and 1900 
m. This species was most common in the 1200 
and 1500 m zones and was distinctly rare at 810 m. 

Diet — On 9 November at 1200 m, a C. ran- 
drianasoloi captured a 1.5-cm-long cricket from 
a leaf about 3 m off the ground, with which it 
flew to the ground. The cricket was beaten on the 
ground for 2 minutes before being eaten; the legs 
were eaten separately. 

Breeding — An individual was observed on 9 
November at 1200 m carrying food, presumably 
for juveniles. On 21 November at 810 m, a 
mixed-species flock contained a group of six C. 
randrianasoloi, including three juveniles. 

Local Name — Simitsy (RNI d'Andohahela, 
parcel 1). 



Nesillas lantzii 

Lantz's Brush- Warbler 

Recent work on the relationships of Nesillas in 
southeastern Madagascar has found that the spiny 
forest and littoral forest form N. (typica) lantzii is 
genetically isolated from the humid forest popu- 
lations (A/. /. typica) and that the two should be 
considered separate species (Schulenberg et al., 
1993). Here we propose the English common 
name Lantz's Brush- Warbler for this species. 

N. lantzii occurs in the spiny forest, east over 
the Col de Ranopiso to Petriky and Tolagnaro 
(fmnh), and then north through the littoral forest 
zone to perhaps Mandena (sight records). Presum- 
ably lantzii is the form occurring west to the Man- 
drare River, although it is not particularly com- 
mon in the forest habitat of Malaza and Bealoka. 
In mid-December this species was not found in 
gallery and spiny forest of parcel 2 of the RNI 
d'Andohahela east of Hazofotsy. 

Lantz's Brush- Warbler is a common bird in in- 
tact to heavily disturbed littoral forest and spiny 
forest from sea level to about 100 m. This species 
can also be found in low scrub and village gar- 
dens (e.g., Tolagnaro) and in relict humid forest 
patches at Pic St. Louis (near Tolagnaro). It typ- 
ically forages close to the ground, skulks in rel- 
atively thick vegetation, and makes its presence 
known by its characteristic rattling contact call. 

Diet — Stomach contents of collected individu- 
als invariably contained insect remains. 

Breeding — None of the specimens we obtained 
in September and October were in or approaching 
breeding condition. 



Weight— Female (1), 15.0 g; male (5), 17.4 ± 
0.8 (16.0-18.0) g; combined (6), 17.2 ± 0.9 
(15.0-18.0) g. 

Soft Part Colors — Bill: maxilla dusky gray 
or black with yellow cutting edge, and mandible 
yellowish gray or light pinkish yellow with darker 
subterminal spot; mouth lining: yellow; legs: dull 
bluish gray; claws: black; iris: medium brown. 



Nesillas typica typica 

Madagascar Brush-Warbler 

The range of the Madagascar Brush- Warbler in 
southeastern Madagascar is from the Marovony 
Forest south through the humid forests of the An- 
osyenne and Vohimena mountains to the Manan- 
tantely Forest from near sea level to 1900 m. It 
also occurs in littoral forest as far south as Man- 
afiafy (Schulenberg et al., 1993). Throughout this 
area it is parapatric with N. lantzii. In the heart of 
the Marosohy Forest the Madagascar Brush-War- 
bler was distinctly more common in scrubby areas 
along river margins and regenerating areas asso- 
ciated with natural landslides than in closed-can- 
opy forest. In parcel 1 of the RNI d'Andohahela, 
this species was absent from humid forest at 440 
m, scarce at 810 m, and abundant at 1200-1900 
m. This pattern has been noted in other humid 
forests (Hawkins et al., in press). In contrast, out- 
side of humid forest, in degraded areas and sec- 
ondary forest, the species is common at much 
lower elevations. 

Diet — The stomachs of three individuals con- 
tained spiders (Araneae), Diplopoda, Blattodea, 
Coleoptera (Nitidulidae, Tenebrionidae), Hemip- 
tera, Homoptera, and Hymenoptera (Apoidea) 
(Goodman & Parrillo, in press). 

Breeding — Two males taken in the Marovony 
Forest in late October had testes up to 12 X 8 
mm. A recent fledgling, still being fed by its par- 
ents, was found in scrub at the edge of the Ana- 
lalava Forest on 5 November. 

Weight— Female (3), 16.0, 16.5, 18.5 g; male 
(6), 18.4 ± 1.9 (17.0-22.0) g; combined (25), 
18.6 ± 2.4 (15.0-26.0) g. 

Soft Part Colors — Bill: maxilla dusky gray 
or black with yellow cutting edge, and mandible 
yellowish gray or light pinkish yellow; mouth lin- 
ing: yellow; legs: pale grayish brown or dull blu- 
ish gray; claws: brown; iris: medium brown and 
grayish brown (subadults). 

Local Name — Parataka. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



69 



Thamnornis chloropetoides 
Thamnornis Warbler 

The eastern boundary of this dry country spe- 
cies extends into the western limit of southeastern 
Madagascar as it is defined herein. Langrand 
(1990) reported this species from the RP de Ber- 
enty. In parcel 2 of the RNI d'Andohahela it was 
rather scarce. Only nine singing birds were locat- 
ed along 5 km of trail; this frequency is much 
lower than that in spiny forest in southwestern 
Madagascar. This species has been found in the 
Euphorbia bush scrub near Lac Anony and seems 
to be more common in coastal areas with appro- 
priate habitat than in inland areas. 



Cisticola cherina 
Madagascar Cisticola 

This species was found in open areas from the 
Marovony Forest south through the low hills of 
the Anosyenne and Vohimena mountains to Petri- 
ky and west to the Mandrare River, from sea level 
to about 1500 m. It was particularly common 
along the coastal plain in low-lying areas of 
heathland, wet grassland, rice paddies, marshes, 
and meadows. Along the lower slopes of the 
mountains this species generally was confined to 
agricultural fields, although we did find it in grass- 
land areas at 800 m near Antseva and at 1500 m 
in a seasonal marsh north of Trafonaomby. West 
of the Anosyenne Mountains it was locally com- 
mon along grass verges and undisturbed grassland 
pastures and in gallery and riverine habitat. In the 
RP de Berenty, it was common in the meadow 
bordering the inland limit of the gallery forest. 

Breeding — An adult female netted at Ankapo- 
ky on 9 October was not in breeding condition. 

Weight — Unsexed (1), 9.7 g. 

Soft Part Colors — Bill: maxilla black with 
gray cutting edge, and mandible gray; legs: gray- 
ish pink; iris: tan brown. 



Dromaeocercus brunneus 
Brown Emutail 

The Brown Emutail is known in southeastern 
Madagascar only from the upper montane zone of 
the RNI d'Andohahela (parcel 1) between 1200 
and 1 800 m. In this zone it was fairly common in 
the understory of forested areas with dense her- 
baceous cover. 



Randia pseudozosterops 
Rand's Warbler 

All of our records of Rand's Warbler in south- 
eastern Madagascar are from the humid forests of 
Marosohy, Marosalohy, and parcel 1 of the RNI 
d'Andohahela. In these forests, which are more or 
less contiguous, it was observed in undisturbed 
forest between 375 and 1600 m. Most of our re- 
cords are of single individuals 20-25 m high in 
closed-canopy forest, often singing from the 
crowns of tall riverside trees. At elevations above 
800 m this species is distinctly less common. On 
several occasions it was observed in mixed-spe- 
cies foraging flocks. This species characteristical- 
ly forages along horizontal branches, hitching 
along the surface of the branch and often peering 
under to check the undersurface of the branches. 

No specimens of this species are known from 
southeastern Madagascar. The previous southern- 
most record was the humid forest in the RNI 
d'Andringitra (Goodman & Putnam, 1996). 

Breeding — On 26 November at 1500 m in par- 
cel 1 of the RNI d'Andohahela, two adults were 
feeding a begging juvenile in the middle of a mul- 
tispecies flock. The juvenile appeared similar in 
plumage pattern and coloration to an adult. 



Newtonia amphichroa 
Dark Newtonia 

In southeastern Madagascar the Dark Newtonia 
inhabits areas of intact humid forest from the Ma- 
rovony Forest south through the Anosyenne and 
Vohimena ranges to the forests of Bezavona and 
Marosohy and portions of parcel 1 of the RNI 
d'Andohahela from above 440 m to about 1900 
m. At this latter site it was absent from forest at 
about 440 m, most common between 810 and 
1500 m, and still fairly common at 1900 m. It was 
not recorded in the transitional forest north of Tra- 
fonaomby between 1500 and 1700 m. A specimen 
was collected 30 km NNW Fort-Dauphin (mnhn). 
This species has not been recorded in the Man- 
antantely Forest or in any littoral forest. 

Juveniles of N. amphichroa are different from 
juveniles of N. brunneicauda in that individuals 
of the former species are distinctly darker and the 
edges of the upper wing coverts are broadly edged 
with reddish brown (fmnh). 

Diet — The stomachs of two specimens collect- 
ed in 1989 contained spiders (Araneae: Saltici- 
dae), beetles (Chrysomelidae, Curculionidae), 



70 



FIELDIANA: ZOOLOGY 






ants (Formicidae), Homoptera, and Orthoptera 
(Goodman & Parrillo, in press). 

Breeding — A fledgling being fed by an adult 
was noted in the Marosohy Forest on 25 Novem- 
ber at about 1 100 m. A similar observation was 
made in parcel 1 of the RNI d'Andohahela on 25 
November at 1500 m. 

Weight— Male (2), 12.0, 12.5 g; combined 
(13), 12.5 ± 1.5 (10.0-15.5) g; subadult (3), 9.5, 
10.5, 10.5 g. 

Soft Part Colors — Bill: black or dark brown; 
mouth lining: yellow; legs and claws: gray; iris: 
golden yellow (adult) and brownish gray (suba- 
dults). 



Newtonia brunneicauda brunneicauda 
Common Newtonia 

This species was the most common and wide- 
spread of the four Newtonia spp. in southeastern 
Madagascar. It was noted in humid, littoral, tran- 
sitional, and spiny forest from Marovony Forest 
south through the Anosyenne and Vohimena rang- 
es to Petriky and west to the Mandrare River. It 
occurs in primary to heavily disturbed forests and 
at the forest edge from near sea level to 1900 m. 
It was recorded in virtually every forest parcel 
visited in the region. The sole exception is the 
Mandena Forest; however, there is a specimen 
taken 7 km north of Tolagnaro (mnhn), which 
would be in or close to the Mandena Forest. This 
species previously has been reported from the Be- 
mangidy Forest (Appert, 1985). Together with 
Neomixis tenella and Nectarinia souimanga, it 
was one of the most common small passerines in 
littoral forest. 

In spiny forest the Common Newtonia was rel- 
atively common in the lower strata of the spiny 
forest, generally below 5 m. In this habitat N. 
brunneicauda is broadly sympatric with N. arch- 
boldi. The Common Newtonia is also found in 
gallery forest, e.g., RP de Berenty. All specimens 
examined from southeastern Madagascar are re- 
ferable to N. b. brunneicauda. 

Diet — The stomachs of collected specimens 
contained spiders (Salticidae), Blattodea, beetles 
(Anobiidae, Buprestidae, Chrysomelidae, Cleri- 
dae, Curculionidae, Elateridae, Scarabaeidae), He- 
miptera, Homoptera (Cicadellidae), Diptera, Hy- 
menoptera (Formicidae), and Lepidoptera (larvae 
and adults) (Goodman & Parrillo, in press). An 
adult taken near Mandena had been feeding on 
insects similar to small cicadas (mnhn). In subarid 



regions this species was regularly observed feed- 
ing in flowering Euphorbia and Sarcostemma. 
The birds may be exploiting insects attracted to 
the nectary glands of the former and to the pun- 
gent, fragrant flowers of the latter. 

Breeding — The vast majority of adult brunnei- 
cauda specimens taken in the area between late 
September and late December were not in breed- 
ing condition. In the RNI d'Andohahela (parcel 
1 ) no evidence of this species breeding was found 
between mid-October and early December. Fledg- 
lings were found at Manafiafy on 10 and 24 Oc- 
tober. In the Ankapoky Forest, however, five 
adults collected during the first portion of October 
were in or approaching breeding condition. Thus, 
it is possible that the breeding of brunneicauda in 
southeastern Madagascar may be much earlier in 
the humid forest areas than in the dry forest areas. 

Weight— Female (8), 10.9 ± 0.7 (9.8-12.0) g; 
male (10), 10.5 ± 1.1 (8.9-12.5) g; combined 
(29), 10.4 ± 1.0 (8.7-12.5) g. 

Soft Part Colors — Bill: black; mouth lining: 
yellow; legs: light pinkish brown or grayish 
brown; claws: gray; iris: pale dull yellow, pale 
cream, golden yellow, or dark brownish gray 
(subadults). One bird with a skull about 25% os- 
sified had a pale yellowish brown iris, and another 
with an 80% ossified skull had a dull yellow iris. 

Local Names — Andreabokia (Berenty), tsi- 
mimitry or tsimimitsy (Marosohy). 



Newtonia archboldi 
Archbold's Newtonia 

In southeastern Madagascar Archbold's New- 
tonia is confined to the spiny forest region, where 
it is relatively common in spiny forest. The east- 
ern limit of this species's range is a few kilome- 
ters west of the eastern boundary of parcel 1 of 
the RNI d'Andohahela, south to the Col de Ran- 
opiso. It occurs in appropriate habitat from the 
Ankapoky Forest west to the Mandrare River. It 
is particularly common in areas dominated by Eu- 
phorbia and Didiereaceae. 

Diet — The stomachs of two collected individ- 
uals contained spiders (Araneae), beetles, Homop- 
tera (Cicadellidae), Isoptera, and Lepidoptera 
(Goodman & Parrillo, in press). 

Breeding — An adult male taken at Ankapoky 
on 9 October had testes 5X4 mm (left) and 4 X 
3 mm (right). 

Weight— Male (2), 7.2, 8.3 g. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



71 



Soft Part Colors — Bill, legs, and claws: 
black; iris: yellowish white. 



Newtonia fanovanae 
Red-tailed Newtonia 

Until 1989 the Red-tailed Newtonia was known 
only from the type specimen collected in 1931 in 
the Fanovana Forest, central eastern Madagascar 
(Gyldenstolpe, 1933). This species was found in 
November and December 1989 and May 1990 in 
the Marosohy Forest between 300 and 1300 m 
(Goodman & Schulenberg, 1991) and in late De- 
cember 1992 in the adjacent forest of parcel 1 of 
the RNI d' Andohahela on the east side of the Col 
d'Ambatomaniha along the Isedro Trail at approx- 
imately 525 m. It also was observed on 22 July 
1992 at 200 m elevation along the Tanatana Trail, 
in the same portion of the reserve (Langrand & 
Sinclair, 1994). In the RNI d' Andohahela (parcel 
1 ) this species was recorded only at 440 m, where 
it was relatively common. 

In southeastern Madagascar N. fanovanae is re- 
stricted to closed canopy forest, and it occurs 
sympatrically with two other species of Newtonia. 
These species show habitat and elevational seg- 
regation: amphichroa is restricted to montane for- 
est and lives in the lower strata of forest between 
810 and 1900 m; brunneicauda is the most wide- 
spread and occurs in the lower and middle strata 
of the forest between sea level and 1950 m; and 
fanovanae occurs in the middle and upper strata 
from 300 to 1375 m (see Goodman & Schulen- 
berg, 1991, for further details). 

Diet — The stomach of one bird was mostly 
empty but contained a few insect parts, presum- 
ably orthopteran. 

Breeding — An adult male specimen collected 
on 8 December had testes 10 X 5 mm (left) and 
8X4 mm (right). 

Weight— Male (1), 12.5 g. 

Soft Part Colors — Bill: maxilla black, and 
mandible horn gray at base merging to light gray 
at tip; mouth lining: horn gray; legs: slate gray; 
claws: light gray; iris: reddish brown. 



Neomixis tenella 
Common Jery 

In southeastern Madagascar the Common Jery 
was common in intact to heavily disturbed littoral 
and humid forest from Marovony south through 



the Anosyenne and Vohimena mountains to Petri- 
ky. In this area it was regularly noted from near 
sea level to 150 m but was rarer at higher eleva- 
tions up to 1 700 m. In the Marosohy Forest it was 
sympatric with N. viridis and N. striatigula. In the 
humid forest of parcel 1 of the RNI d' Andohahela 
this species was most common at 440 m and 810 
m, rarer at 1200 m, and completely absent above 
1500 m. However, it reappeared in transitional 
forest north of Trafonaomby, at elevations from 
1500 to 1700 m. This species often was found in 
the ecotone of secondary littoral forest and heath- 
land or grassland areas, where it was one of the 
most common birds, e.g., near the forests of Man- 
afiafy, Mandena, Itapera, and Petriky. It also was 
a garden bird of coastal towns and villages, in- 
cluding Tolagnaro. In this region the local sub- 
species is N. t. orientalis, as is a specimen taken 
near Lac Anony (pbzt). 

The Common Jery was distinctly less common 
in the spiny forest region west of the Anosyenne 
Mountains. In this area it was found in spiny for- 
est, often sympatric with N. striatigula, and in gal- 
lery forest, for example along the Mandrare River. 
The form occurring in this region is presumably 
N. t. debilis. 

This species was often noted foraging in small 
groups, both in single-species flocks of up to 15 
individuals and in mixed-species flocks. 

The juveniles observed in parcel 2 of the RNI 
d' Andohahela were brighter than the adults, with 
clean pale yellow breasts lacking the olive suf- 
fusion and streaking of the adults, orange gapes, 
and clean gray napes; they thus resembled adults 
of the race N. t. tenella, except that they had dark 
brown instead of pale irides. The adults had clear 
yellow throats and were heavily tinged olive on 
the sides of the breasts and narrowly streaked dark 
olive in the center. The crown was greenish yel- 
low, the back was fairly bright greenish, and the 
nape was dull greenish gray. The legs were bright 
pale pink and the lower mandible of the bill was 
pale pinkish orange and the upper mandible was 
dark horn. 

Diet — The stomachs of collected individuals 
contained arthropods, including spiders (Araneae: 
Salticidae), Blattodea, beetles (Chrysomelidae, 
Coccinellidae, Curculionidae, Scarabaeidae, Sco- 
lytidae), Diptera (Nematocera), Homoptera, Hy- 
menoptera (Formicidae), Orthoptera (Gryllidae), 
and Lepidoptera (larvae and adults) (Goodman & 
Parrillo, in press). This species has been observed 
in Tolagnaro feeding at domestic Hibiscus flowers 
and at Bealoka at Capparis flowers. It is often 



72 



FIELDIANA: ZOOLOGY 









seen searching for insect larvae on the undersides 
of leaves, but it also will take adult insects (i.e., 
lacewings, which are easy prey). In dry areas they 
also visit the flowering heads of sisal and appar- 
ently prefer to search for food in vines and trees 
with compound leaves, Acacia rovumae being the 
most favored. 

Breeding — Numerous adults collected in litto- 
ral forest areas between mid-September and late 
October had enlarged gonads, including males 
with testes up to 9 x 6 mm and females with 
ovaries 7X2 mm and ova up to 1 mm in diam- 
eter. Fledglings and subadults were noticed at 
Manafiafy in mid- to late October and at Anala- 
lava on 9 November. An adult pair in parcel 2 of 
the RNI d'Andohahela on 12 December was ac- 
companied by two well-grown juveniles. Males in 
breeding condition lacked brood patches. In sub- 
arid areas active nests have been found in Novem- 
ber and fledglings have been found in mid-De- 
cember. 

Weight— Female (2), 7.0, 8.5 g; male (11), 7.2 
± 0.5 (6.8-8.2) g; combined (21), 7.3 ± 0.5 (6.8- 
8.5) g. 

Soft Part Colors — Bill: maxilla dark gray to 
black, and mandible dull pink sometimes with 
dusky tip and yellowish pink in fledglings; legs: 
light orangish brown; claws: gray or blackish 
gray; iris: light to medium brown. 

Local Names — tsimimitraka (Marosohy), tsim- 
itsy (Berenty, Bealoka), zea-zea (Manafiafy), 
which is generic for Neomixis or perhaps small 
bird. 



Neomixis viridis 
Green Jery 

The Green Jery was uncommon and confined 
to intact humid forest parcels. We observed this 
species in the Marosohy Forest and parcel 1 of 
the RNI d'Andohahela between 350 and 1950 m 
and in the Bezavona Forest at about 75 m. In the 
RNI d'Andohahela it was present at all elevations 
from 440 to 1950 m but was most common at 810 
and 1200 m and was distinctly scarce at 440 m. 
Langrand (1990) reported this species as occur- 
ring as far south as Tolagnaro. No specimens from 
the region are available, but presumably the local 
form is N. v. viridis. 

Breeding — On 2 November at 810 m in parcel 
1 of RNI d'Andohahela, a group of three birds 
included a food-begging juvenile. The plumage of 
the juvenile was similar to that of the adult. 



Neomixis striatigula pallidior and Neomixis 
striatigula [striatigula] 
Stripe-throated Jery 

The Stripe-throated Jery was fairly common in 
intact humid forest at lower elevations (although 
recorded in montane forest at a few places) and 
more common in spiny forest. We found it in the 
Marovony Forest at approximately 50 m and in 
the Marosohy Forest and RNI d'Andohahela (par- 
cel 1) between 440 and 1500 m. During an in- 
ventory of the latter forest, this species was pres- 
ent in all zones from 440 m to 1500 m but was 
much scarcer at and above 1200 m. Only odd in- 
dividuals (easily detectable by their loud and per- 
sistent song) were found at 1200 and 1500 m. It 
also has been observed at about 1150 m in the 
forest east of Vohibaka in the extreme northwest- 
ern corner of the RNI d'Andohahela, and it was 
common in transitional forest at 1500-1700 m, 
north of Trafonaomby. 

Specimens are not available from the humid 
forest of the region, but we presume that this pop- 
ulation is of N. s. striatigula. The distribution and 
southern limits of N. s. sclateri and N. s. striati- 
gula are poorly defined (Salomonsen, 1934; Lan- 
grand, 1990). 

In the spiny forest region it is common, seem- 
ingly more so than in humid forest or at least 
more conspicuous, often singing in the tallest sub- 
strate available in relatively open places. In this 
region it has been recorded at a wide range of 
spiny forest sites, e.g., near Ankapoky and Ha- 
zofotsy, near gallery forest and Euphorbia bush 
at Bealoka and Berenty, and along the Mandrare 
River. The subspecies occurring in the spiny for- 
est is N. s. pallidior (fmnh). The individuals oc- 
curring in the transitional forest, being isolated 
from populations in humid forest, may belong to 
the subdesert subspecies N. s. pallidior. 

Diet — The stomachs of two individuals col- 
lected in the Ankapoky Forest contained beetles 
(Curculionidae), Hemiptera, ants (Formicidae), 
and crickets (Gryllidae) (Goodman & Parrillo, in 
press). 

Breeding — Birds collected at Ankapoky in 
mid-October were in breeding condition. Adult 
males with large testes lacked brood patches. 

Weight— Combined (6), 8.6 ± 0.6 (7.8-9.5) g. 

Soft Part Colors — Bill: maxilla black or 
brownish black, and mandible gray at base merg- 
ing to black tip; legs: grayish pink; claws: black; 
iris: orangish red or orangish brown. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



73 



Hartertula flavoviridis 
Wedge-tailed Jery 

In southeastern Madagascar the Wedge-tailed 
Jery was only recorded in the humid forests of 
Marosohy at about 425 m, in Bezavona at about 
85 m, and in the RNI d'Andohahela (parcel 1) 
between 440 and 1 500 m. At the latter site it was 
seen once at 440 m and was most common from 
810 to 1200 m. In the Marosohy Forest this spe- 
cies was noted several times in multispecies flocks 
with Phyllastrephus madagascariensis and P. zos- 
terops. It generally foraged and moved quickly in 
the lower and middle strata of closed-canopy for- 
est. 

Diet — The stomach of an individual obtained 
in the Marosohy Forest contained Dermaptera 
(Goodman & Parrillo, in press). 

Breeding — On 31 October at 810 m in parcel 
1 of the RNI d'Andohahela, a family group con- 
sisting of a pair and two juveniles was seen. The 
juveniles resembled the adults except that the 
heads were uniform dull olive, the tails were 
short, and the underparts were dull green yellow. 
The fleshy gape was conspicuously yellow. An- 
other group on 21 November at 1200 m consisted 
of two adults and three juveniles. These juveniles 
were older than those found on 31 October and 
were similar to the adults except that they had 
olive, not gray, ear coverts. A female collected in 
the Marosohy Forest on 1 1 December had 90% 
of the skull ossified and a nonenlarged ovary. 

Weight — Female (1), 9.4 g; combined (4), 9.4 
± 0.3 (9.2-9.8) g. 

Soft Part Colors — Bill: maxilla black with 
gray cutting edge, and mandible gray with black 
spot as base and occasionally with black tip; legs: 
horn gray; feet: dull yellow or yellowish green; 
iris: brown. 

Local Name — Tsimimitrala (Marosohy). 



Monarchidae 

Pseudobias wardi 
Ward's Flycatcher 

The only locality in southeastern Madagascar 
in which Ward's Flycatcher has been noted has 
been in the contiguous forests of Marosohy, Ma- 
rosalohy, and the RNI d'Andohahela (parcel 1). 
In this area it has an elevational range between 
440 and 1500 m but was most common between 
810 and 1200 m. This species was often noted 



perched in relatively open forested areas, gener- 
ally at the edge of ravines, from where it would 
sally out and capture insects on the wing. Speci- 
mens in mnhn were collected 30 km NNW Fort- 
Dauphin. 

Diet — The stomachs of two individuals col- 
lected 30 km NNW Fort-Dauphin on 28 May con- 
tained exclusively insect remains (mnhn). 

Breeding — An adult was observed carrying 
food, presumably for a juvenile, on 2 November 
at 810 m in parcel 1 of the RNI d'Andohahela. 
The testes of one of the specimens (mnhn) men- 
tioned above were 1.7 X 0.7 mm (left) and 1.6 X 
0.7 mm (right). 



Terpsiphone mutata mutata 

Madagascar Paradise Flycatcher 

In southeastern Madagascar the Madagascar 
Paradise Flycatcher was noted from the Marovony 
Forest south through the Anosyenne and Vohi- 
mena mountains to Petriky and west across the 
spiny forest region to the Mandrare River, from 
near sea level to 1800 m. Although this species 
was common in a variety of forest habitats, it ap- 
peared to achieve its highest densities in second- 
ary littoral or humid forest, such as portions of 
the Manafiafy Forest. It was recorded in virtually 
every littoral and humid forest site visited. In par- 
cel 1 of the RNI d'Andohahela, this species was 
recorded at all elevations sampled below 1 800 m, 
although it was most common between 440 and 
810 m and rare at 1500 m. On a few occasions 
individuals were observed in gardens on the out- 
skirts of Tolagnaro. A specimen was collected in 
1756 near Fort-Dauphin (Stresemann, 1952). 

In the spiny forest region it was distinctly less 
common than in more humid areas to the east, al- 
though it is broadly distributed across the spiny 
forest. In areas with gallery forest, such as along 
the Mandrare River, however, populations are 
dense. In the RP de Berenty a Terpsiphone was 
caught in the large webs of the colonial Nephila 
spider. 

Adult males in definitive plumage exhibit con- 
siderable variation in coloration: the tail streamers 
and breast may be white or rufous, the throat and 
rectrices (other than the central pair) may be black 
or rufous, and the back may be black, white, or 
rufous. Certain combinations of plumage patterns 
occur sufficiently frequently that these have been 
interpreted as representing four discrete color 
phases. On the basis of the geographic distribution 



74 



FIELDIANA: ZOOLOGY 



of the frequency of color morphs, two subspecies 
have been recognized (Salomonsen, 1933a, b). We 
suspect that plumage variation in fact is more com- 
plex than Salomonsen realized and that the plum- 
age sequences of this species are not well under- 
stood. Until the sequences of plumages and genet- 
ics of polymorphisms are better understood, we re- 
frain from recognizing subspecies on Madagascar. 

Yamagishi et al. (1992) noted that all fledglings 
(unsexed) in four nests, with both "white morph" 
(three nests) and "rufous morph" (one nest) 
males in attendance, were all rufous. From this 
they suggested that this species has delayed plum- 
age maturation, and that "white morph" males 
are older than "rufous morph" birds. We have 
independent evidence of delayed plumage matu- 
ration in males (see below), but we are less con- 
vinced by their suggestion that the definitive "ru- 
fous morph" is a plumage that precedes acquisi- 
tion of "white morph" plumage. It is surprising 
that such basic questions remain unanswered, be- 
cause many of these questions could be addressed 
fairly easily with color-marked birds. R. Mulder 
is currently conducting genetic studies of a color- 
banded population in the RP de Berenty area, and 
over the next few seasons some of these questions 
should be answered. 

We collected several rufous males in October 
and November that were in a plumage identical 
to that of the females, with brown wing coverts, 
no or very short rufous tail streamers, and pale 
reddish brown underparts. These males had fully 
ossified skulls and enlarged testes (up to 5 X 10 
mm). From the degree of skull ossification, these 
males clearly were not recently fledged birds, and 
on the basis of the enlarged gonads, they may 
have been in reproductive condition. These spec- 
imens strongly suggest that this species has de- 
layed plumage maturation. 

Diet — Stomach contents of collected individu- 
als included spiders (Araneae), Orthoptera, beetles 
(Cuculionidae, Elateridae, Scarabaeidae), Homop- 
tera, and Hymenoptera. On the basis of stomach 
contents and direct observations, the diet of this 
species is exclusively insects and includes butter- 
flies, moths, and lacewings. This species has been 
observed following Coua gigas and feeding on 
insects disturbed by them. In gallery forest Terp- 
siphone was most often seen in Rinorea greveana 
and Tamarindus indicus, which they use as perch- 
es from which to hawk insects. 

Breeding — Most of the adults collected in the 
littoral forests of Manafiafy, Mandena, and Petri- 
ky from mid-September to early October appeared 



to be just entering the breeding season. By mid- 
October there was a noticeable enlargement of the 
gonads and nest building activity was observed, 
although copulations were observed as early as 6 
October. From late November to mid-December 
this species was found in the Marosohy Forest in 
various stages of breeding: adult males were hold- 
ing and defending territories, a female had a par- 
tially shelled egg in the oviduct, adults were ob- 
served feeding fledglings, and subadults with par- 
tially ossified skulls were collected. Males in 
breeding condition lacked brood patches. On 23 
October at 810 m in parcel 1 of the RNI 
d'Andohahela, a female was incubating a nest sit- 
uated 6 m up in a tree; nests are generally within 
2 m of the ground. In the same forest at 1 200 m 
on 9 November, a male and female were feeding 
a single recently fledged juvenile. 

In the dry forests of Malaza and Bealoka the 
open-cup nests of Terpsiphone, often placed low 
to the ground, were relatively easy to find. The 
highest density of breeding birds noted in this re- 
gion was in reserve 3 of the RP de Berenty, which 
is dominated by dense stands of Pithecellobium 
duke. Nests were also placed in thorny shrubs 
such as Capparis seppiaria or Azima tetracantha 
and in nonspiny saplings, e.g., Rinoria greveana. 
In this region, the first half of October is a period 
of nest building, and by mid-November fledglings 
are seen, although in some years the breeding sea- 
son is earlier and fledglings are noted in early Oc- 
tober. 

Weight— Female (11), 13.9 ± 1.4(12.0-17.0) 
g; male (23), 13.8 ± 1.0 (12.0-15.5) g; combined 
(66), 14.0 ±1.1 (12.0-17.0) g. 

Soft Part Colors — Bill: bright bluish gray or 
with black tip, and sometimes mandible complete- 
ly black; mouth lining: yellow or greenish yellow; 
legs: gray or dark bluish gray; claws: black; iris: 
dark brown; orbital ring: bright purplish blue or 
bright blue in males in breeding plumage and less 
developed and often less brilliant in females. 
Fledglings — Bill: brown with bright yellow gape; 
mouth lining: bright yellow; legs and claws: 
creamy gray. 

Local Names — Angetry (Manombo), rengetry 
(Manafiafy, Marosohy), rimaly (Berenty, Bealoka). 

Timaliidae 

Oxylabes madagascariensis 
White-throated Oxylabes 

The White-throated Oxylabes was relatively 
common in the humid forest sites of Marovony, 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



75 



Marosohy, RNI d'Andohahela (parcel 1), Beza- 
vona, and Manantantely between 50 and 1900 m. 
In almost all cases it was noticed or heard calling 
from within 1-2 m of the ground and was often 
difficult to observe because of its skulking habits. 
In the RNI d'Andohahela (parcel 1) this species 
was observed in all elevational zones between 440 
and 1900 m, although it was most common at 810 
and 1200 m and scarce at the highest elevations. 
This species has been collected 30 km NNW Fort- 
Dauphin (mnhn) and near Eminiminy (smf). A 
specimen was taken in 1756 near Fort-Dauphin 
(Stresemann, 1952). This species was not record- 
ed at any littoral forest site in the region. 

On 28 September in the Manantantely Forest, 
at approximately 75 m and 16h00, a diurnal for- 
est-dwelling rodent, Nesomys audeberti, was ob- 
served on the forest floor digging in the leaf liter. 
On the ground near the rodent were two Oxylabes 
madagascariensis, three Phyllastrephus zoster- 
ops, and at least two unidentified species of birds. 
The rodent soon became aware of human pres- 
ence and fled further into the forest. Whether the 
interaction between birds and rodent was a coin- 
cidence, whether the birds were momentarily at- 
tracted to the rodent's activity, or whether they 
were following it to catch prey flushed by the ro- 
dent remains unclear. 

Diet— A bird collected 30 km NNW Fort-Dau- 
phin had insect remains in its stomach (mnhn). 
Stomach contents of individuals collected in 1989 
and 1990 contained spiders (Araneae), beetles 
(Scarabaeidae), Homoptera, and Orthoptera 
(Goodman & Parrillo, in press). 

Breeding — On 28 September in the Manantan- 
tely Forest a pair of adults was noted with at least 
two fledglings. On 3 November in the Marovony 
Forest an adult female was netted with two fledg- 
lings. Breeding records from the RNI d'Andohahela 
(parcel 1) include an adult carrying food, presum- 
ably to a nest, on 30 October at 810 m and seven 
observations of adults with fledglings, between 28 
September and 22 November; six of the seven ob- 
servations were of adults with two fledglings and 
in one observation there was a single fledgling. 
An open cup-shaped nest was located and pho- 
tographed on 2 December at 1340 m in the An- 
tanandava Forest of the RNI d'Andohahela (B. 
Fisher, pers. comm.). The nest was 1.2 m off the 
ground near a rocky outcrop and contained two 
eggs. 

Weight— Combined (9), 23.3 ± 2.8 (18.0- 
27.5) g; subadult (2), 21.5, 23.0 g. 

Soft Part Colors — Bill: maxilla brownish 



black to black with cold gray cutting edge, and 
mandible variable from cold gray to black at base 
with gray along most of length; legs: dull brown 
or slate black; iris: brown. Fledglings — Bill: max- 
illa brownish black with dull yellow tip, and man- 
dible dull yellow with brown spot near tip; legs: 
brown; iris: dark tan. 



Crossleyia xanthophrys 
Yellow-browed Babbler 

In southeastern Madagascar the Yellow-browed 
Babbler was only noted in the Marosohy Forest 
between 425 and 850 m and in the RNI 
d'Andohahela (parcel 1) between 800 and 1800 
m. This species was generally observed on moist 
ground in closed-canopy forest, often in areas 
with dense stands of climbing bamboo. 

Juveniles are duller and darker in plumage col- 
oration than are the adults, particularly the super- 
cilium, breast, and underparts. Adults had all pale 
pink bills, whereas the bills of juveniles had 
darker culmens and tips. 

Diet — The stomach contents of a collected bird 
consisted exclusively of small insects, probably 
beetles. 

Breeding — The following information con- 
cerns observations in parcel 1 of the RNI 
d'Andohahela. Between 13 and 19 November 
pairs of adults were observed at 1 200 m and 1 500 
m with single juveniles or groups of two juve- 
niles. A male obtained on 9 November at the same 
elevation had testes 5X4 mm (left) and 4X3 
mm (right) and no brood patch. On 25 November 
at 1500 m, an adult was feeding at least two new- 
ly fledged juveniles. A pair was observed building 
a nest between 21 and 24 November, at 1500 m. 
The base of the nest was large and bulky, about 
1 5 cm deep, and was made of bamboo leaves and 
branches. This may have been a natural accumu- 
lation of vegetable material on which the nest was 
built. The nest itself was a deep cup, 6-8 cm 
across, lined with bamboo leaves and moss and 
about 2 m above the ground. Two adult birds were 
observed adding bamboo leaves and moss to the 
nest. When an adult sat in the nest, the bird was 
almost hidden, with only the top of the head and 
the end of the tail protruding. 

Weight— Combined (4), 22.4 ± 4.1 (19.5- 
28.5) g. 

Soft Part Colors — Bill: variable from ivory 
pink with brown around nasal operculum to com- 



76 



FIELDIANA: ZOOLOGY 



pletely grayish pink; legs: pinkish gray to dull 
gray; iris: dark brown. 



Mystacornis crossleyi 
Crossley's Babbler 

In southeastern Madagascar Crossley's Babbler 
was restricted to the humid forests of Marovony, 
Analalava, Marosohy, RNI d'Andohahela (parcel 
1), Bezavona, and Manantantely between 50 and 
1500 m. We have no records of it in littoral forest. 
This species apparently is confined to areas of in- 
tact humid forest with closed canopy and open 
understory. It frequently was noted foraging on or 
next to fallen trunks or rotten trees, where the 
babbler would probe mosses and epiphytes with 
its bill. 

The juvenile plumage coloration is dull brown- 
ish gray above and warmer brown below, with a 
whitish moustache and small spot behind the eye, 
as in the adult. 

Diet — Stomachs of three individuals contained 
spiders (Araneae: Salticidae), Blattodea, Dermap- 
tera, Hemiptera, Homoptera, Orthoptera (Acridi- 
dae), and Hymenoptera (Formicidae) (Goodman 
& Parrillo, in press). 

Breeding — On 13 November at 1200 m in par- 
cel 1 of the RNI d'Andohahela, a male was feed- 
ing a well-grown juvenile. On 7 November in the 
Analalava Forest a subadult with a partially os- 
sified skull was collected. A female with an en- 
larged ovary and two corpora lutea was collected 
in the Bezavona Forest on 27 December. A male 
taken on 3 December in the Marosohy Forest had 
testes 5X3 mm and no brood patch. 

Weight— Female (1), 23.5 g; male (2), 24.0, 
28.5 g; combined (6), 24.6 ± 2.7 (21.0-28.5) g; 
subadult (1), 24.5 g. 

Soft Part Colors — Bill: maxilla black with 
grayish blue cutting edge, and mandible grayish 
blue; mouth lining: yellow; legs: grayish pink or 
pale gray; claws: grayish white or gray; iris: 
brown. 

Local Name — Fatsatsatry (Marosohy). 

Nectariniidae 

Nectarinia souimanga souimanga and 
Nectarinia souimanga apolis 
Souimanga Sunbird 

The Souimanga Sunbird is one of the most 
common and widespread birds in southeastern 



Madagascar from the Marovony Forest south 
along the coast and through the Anosyenne and 
Vohimena mountains to the southern coast and 
west across the spiny forest to the Mandrare Riv- 
er, from near sea level to 1900 m. In the humid 
forest of the RNI d'Andohahela (parcel 1) this 
species was recorded along the complete eleva- 
tional transect, although it was most frequent be- 
tween 810 and 1900 m and seemed markedly less 
common at 440 m. This species frequents areas 
of intact and secondary littoral, humid, spiny, and 
gallery forests, heavily degraded habitats, and 
gardens and agricultural areas. It was encountered 
at every forest visited, regardless of condition. In 
and at the edge of littoral forest this species was 
often abundant. Throughout most of the region it 
was sympatric with N. notata. 

Specimens collected at Ankapoky are referable 
to N. s. apolis, whereas specimens from other lo- 
calities east of the Anosyenne Mountains are of 
N. s. souimanga. The only exception is material 
from Petriky, which appears to be intermediate 
between N. s. souimanga and apolis. 

Diet — A bird taken near Mandena had spider 
remains in its stomach (mnhn). The vast majority 
of stomachs of specimens we examined contained 
spiders (Araneae), although a few individuals 
were found of Hemiptera, Homoptera (Cicadelli- 
dae), Coleoptera (Elateridae), Orthoptera, and Hy- 
menoptera (Apoidea, Formicidae) (Goodman & 
Parrillo, in press). 

Flower nectar would probably be indiscernible 
in the stomach remains. Flowering plants that A/. 
souimanga is known to visit include Citrus, Mal- 
va, Hibiscus, Mascarenhasia, Euphorbia tirucal- 
li, Agave rigida, Opuntia, Delonix regia, Tamar- 
indus indica, Dicoma, Acacia farnessiana, A. 
sakalava, A. rovumae, Aibizia polyphylla, Kalan- 
choe beharensis, Aloe vaombe, A. divaricata, 
Cynanchum spp., Cordia spp., Sarcostemma de- 
corsei, Capparis chrysomeia, Maeura filiformis, 
Eucalyptus citriodora, and Fernandoa madagas- 
cariensis. In most cases N. souimanga feed on 
nectar of these plants, but it also takes advantage 
of blossoms, such as those of Asclepiadaceae 
and Fabaceae, to catch small insects that are at- 
tracted to the flowers. Although flowers of over 
25 species of plants provide N. souimanga with 
food in the gallery forests of Berenty and Bealo- 
ka, Tamarindus flowers are a dominant food re- 
source between the months of January and June. 
Also, the sunbirds' territoriality over patches of 
Aloe indicates guarding of the flowers. In nu- 
merous cases the larger and longer-billed N. no- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



77 



tata was observed frequenting the same plants as 
N. souimanga, and there was interspecific ag- 
gression. N. souimanga also visits Agave flowers 
to catch insects and drink sap from holes gouged 
in the stalks by Lemur catta. 

Although N. souimanga is presumed to be a 
year-round resident in southeastern Madagascar, 
there are seasonal influxes of N. souimanga in 
gallery and spiny forest that appear to be corre- 
lated with the flowering of certain plants. During 
September and October at Hazofotsy, Aloe divar- 
icata is in flower, and considerable numbers (per- 
haps hundreds) of N. souimanga may occur in an 
area of about 2 ha. Likewise in the botanical gar- 
den of RP de Berenty, flowers of A. vaombe, 
which blossom in July, have the same effect. Both 
Fernandoa madagascariensis and Kalanchoe be- 
harensis are among the most commonly visited 
plants, and in July and August, when in flower, 
there is a noticeable increase in the number of 
sunbirds. 

At the summit of Trafonaomby ( 1 960 m) on 26 
November, two female N. souimanga were appar- 
ently disputing feeding access to a dense clump 
of Bakerella flowers. One female successfully de- 
fended the patch by perching next to it and flap- 
ping her wings slowly while opening and closing 
her bill in the rhythm of the wingbeats. She did 
not call while displaying. On 1 December, two 
female N. souimanga disputed the patch with a 
male Neodrepanis hypoxantha. 

Breeding — Numerous individuals taken 
throughout the region and in various types of hab- 
itats between late September and late December 
had enlarged gonads and appeared to be breeding. 
Evidence of nesting was found at several littoral 
sites in October. A nest with two eggs was found 
at Petriky on 2 October. On 8 October a female 
that had a partially shelled egg in the oviduct was 
collected as she was building a nest. Fledglings 
were observed or collected in the months of Sep- 
tember, October, and November. Between 1 and 6 
November in the RNI d'Andohahela at 810 m, a 
male and a female were feeding at least two nest- 
lings in a spherical or ovoid nest placed 1.5 m off 
the ground in an area of dense vegetation. Males 
lacked brood patches. 

Weight — Nectarinia s. souimanga female 
(16), 6.9 ± 0.6 (5.5-7.6) g; male (16), 7.6 ± 0.8 
(5.5-8.5) g; combined (43), 7.2 ± 0.7 (5.5-8.5) 
g. N. s. apolis female (2), 6.0, 6.0 g; male (3), 
6.2, 6.7, 7.0 g; combined (7), 6.2 ± 0.5 (5.2-7.0) g. 

Soft Part Colors — Bill: black and in sub- 
adults distinct yellow fleshy gape; legs and claws: 



black or dark brown and in fledglings gray; iris: 
dark brown. No difference was found in soft part 
colors between N. s. souimanga and N. s. apolis. 
Local Names — Bitraky, generic for Nectarinia 
spp. (Manombo); soy (RNI d'Andohahela, parcel 
1); soy kely (Marosohy); soy manga (Manafiafy); 
tsiksoysoy (Berenty, Bealoka, Hazofotsy). 



Nectarinia notata notata 
Long-billed Green Sunbird 

In southeastern Madagascar the Long-billed 
Green Sunbird is found from the Marovony Forest 
south along the coast and through the Anosyenne 
and Vohimena mountains to Petriky and west 
across the spiny forest to the Mandrare River be- 
tween near sea level and 1200 m. This species 
was relatively common in littoral and humid for- 
ests in both pristine and heavily disturbed condi- 
tion, but was distinctly less common in the spiny 
forest area. It was noted on occasion in gardens 
of Tolagnaro and Manambaro. In general this spe- 
cies was broadly sympatric with N. souimanga but 
was less common. 

In dry and spiny forest areas N. notata ap- 
peared to be seasonally migratory and was often 
absent for long periods. Between July and Sep- 
tember it arrived in the region, usually in low 
numbers. During this period N. notata exhibits 
territorial behavior, i.e., guarding the flowers of 
ephemerally available food plants to the point of 
exhaustion, particularly to thwart the visits of 
conspecifics and N. souimanga. For example, on 
20 August individuals of N. notata were defend- 
ing different flowering Fernandoa trees in the 
Malaza Forest. One N. notata was observed 
chasing five different N. souimanga from its 
patch. The notata returned to the site and would 
occasionally feed at the tree's flowers. Ten days 
later, no notata were observed in the immediate 
area of the Fernandoa trees. Near Hazofotsy this 
species also has been observed between July and 
February. 

Diet — Two birds collected near Mandena had 
spider debris in their stomachs (mnhn). The ma- 
jority of specimens collected in 1989 and 1990 
had arthropod remains in their stomachs, includ- 
ing spiders (Araneae), beetles (unidentified adults 
and larvae), and ants (Formicidae) (Goodman & 
Parrillo, in press). The exception was an adult fe- 
male taken in the Analalava Forest on 9 Novem- 
ber that had a quantity of small seeds in her stom- 
ach. In the Ankapoky Forest we saw this species 



78 



FIELDIANA: ZOOLOGY 









feeding on the flowers of Opuntia cactus. Near 
Berenty and Hazofotsy the favored flowers are 
various leguminous plants, Aloe divaricata, A. 
vaombe, Fernandoa madagascariensis, and Agave 
rigida. 

Breeding — Several specimens taken in October 
and November had enlarged gonads, including 
males with testes up to 8 x 6 mm and females 
with enlarged ovaries and ova up to 6 mm in di- 
ameter. On 14 September at Mandena a fledgling 
was observed being fed by an adult male. Birds 
in juvenile plumage and with partially ossified 
skulls were obtained at Manafiafy on 18 October. 

Weight— Female (5), 13.8 ± 1.7 (12.0-16.0) 
g; male (9), 15.4 ± 1.2 (13.8-17.0) g; combined 
(17), 14.8 ± 1.4 (12.0-17.0) g. 

Soft Part Colors — Bill, legs, and claws: 
black; iris: dark brown. 

Local Names — Bita (Marovony), katia (Man- 
afiafy), soy lehibe (Marosohy). 



Zosteropidae 

Zosterops maderaspatana maderaspatana 
Madagascar White-eye 

The Madagascar White-eye was found in hu- 
mid forest from the Marovony Forest south 
through the Anosyenne and Vohimena moun- 
tains to the Manantantely Forest. It was noted 
in forest interior and forest edge and in dis- 
turbed areas. Mandena was the only littoral for- 
est in which it was found, where the only record 
was a single mist-net capture. This species's 
elevational range is from near sea level to 1950 
m. A specimen dated 1756 was collected near 
Fort-Dauphin (Stresemann, 1952). In general 
this species was uncommon in dry forest, al- 
though it was seasonally common in the RP de 
Berenty and in parcels 2 and 3 of the RNI 
d'Andohahela. An individual was collected by 
Bluntschli near Amboasary-Sud in early No- 
vember (smf). 

The Madagascar White-eye is one of the most 
regular members of mixed-species flocks. In hu- 
mid forest of Marosohy above 425 m, 53% of 
the observed mixed-species flocks contained 
this species. This percentage is similar (over 
60%) to that for flocks containing Zosterops re- 
ported from the RS d'Analamazaotra (P6rinet) 
by Eguchi et al. (1992), who considered Zoster- 
ops to be a nuclear species in mixed-species 
flocks. In some cases considerable numbers are 



observed. For example, on 25 May, along the 
Tanatana Trail in parcel 1 of the RNI 
d'Andohahela at about 200 m, a mixed-species 
flock was observed that contained two Coraci- 
na, at least one Hypsipetes, two Newtonia brun- 
neicauda, two Terpsiphone, 25 Zosterops, one 
Leptopterus viridis, two Cyanolanius, and two 
Dicrurus (OL & SOC). 

Diet — Individuals taken in the general Tolag- 
naro area (mnhn) had a variety of seeds in their 
stomachs, including those from a plant locally 
known as arongana. The stomach contents of 
individuals we collected contained gastropods, 
spiders (Araneae), Blattodea (ootheca), beetles 
(Curculionidae), Homoptera, Diptera, Hyme- 
noptera, and small seeds (Goodman & Parrillo, 
in press). In the Marosohy Forest at 900 m a 
flock of over 15 Madagascar White-eyes was 
observed feeding on the fruits of a Macaranga. 
In dry areas, Zosterops is a relatively common 
visitor to sisal flowers and flowers of Tamar- 
indus indica, Rinoria greveana, Capparis chry- 
someia, Crateva excelsa, and Cordia ronnii, to 
fruits of Phyllanthus seyrigii, and to leaf shoots 
of Acacia rovumae. Visits to flowers may not 
always be for nectar, and this species may con- 
sume pollen and stamens of Crateva and Aca- 
cia. 

Breeding — None of the eight specimens col- 
lected by Milon in February and May near To- 
lagnaro (mnhn) had enlarged gonads. Several 
individuals obtained in more recent years in No- 
vember and December were in or approaching 
breeding condition (fmnh). For example, a fe- 
male taken on 7 November in the Analalava 
Forest (weight = 13.5 g) had a partially shelled 
egg (weight = 2.5 g) in the oviduct and no cor- 
pus luteum, and the largest ovarian follicles 
were 7, 5, and 3 mm. Males with large testes 
lacked brood patches. On 28 November in the 
Marosohy Forest a pair of birds was feeding two 
fledglings. On 21 November at 1200 m in parcel 
1 of the RNI d'Andohahela a mixed-species 
flock contained at least 10 Zosterops, with three 
or four juveniles. 

Weight— Female (4), 11.3 ± 2.1 (8.5-11.5) g; 
male (5), 10.4 ± 0.9 (9.2-1 1.5) g; combined (34), 
10.7 ± 1.4 (8.5-13.5) g. 

Soft Part Colors — Bill: maxilla black, and 
mandible black along distal 50-75% with gray 
base; legs: dark gray; claws: gray; iris: dark 
brown. 

Local Names — Angoiky (Marosohy), bemaso 
(Berenty), siay (Manombo). 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



79 



Vangidae 

Calicalicus madagascariensis 
Red-tailed Vanga 

The Red-tailed Vanga was relatively common 
in closed-canopy humid forests from Marovony 
south through the Anosyenne and Vohimena rang- 
es to Manantantely. We recorded this species be- 
tween 50 and 1600 m. In parcel 1 of the RNI 
d'Andohahela it was recorded between 440 and 
1500 m but was most common at 440 and 810 m 
and much scarcer above 1200 m. Appert (1985) 
reported this species from Bemangidy. There are 
also specimens (mnhn) taken 30 km NNW Fort- 
Dauphin. There is a specimen dated 1756 from 
"Fort-Dauphin," which is also the restricted type 
locality of this species (Stresemann, 1952; Rand, 
1960). This species was noted in both pristine and 
heavily degraded humid forest. We did not record 
this species in littoral forests of southeastern Mad- 
agascar. In the 1960s or 1970s G. Randrianasolo 
observed Calicalicus near the Station Forestiere 
de Mandena, a littoral forest site. The station is 
less than 2 km from the edge of the humid forest; 
however, it is not clear if this species was ob- 
served at Mandena proper. It is generally absent 
from the spiny forest of southeastern Madagascar, 
although it has been recorded in the RP de Ber- 
enty (Langrand, 1990; Nagata et al., 1992). 

The Red-tailed Vanga was regularly noted in 
mixed-species flocks occurring in humid forests; 
it was a member of over 70% of plurispecific 
flocks observed. There is circumstantial evidence 
that this species may play a central role in the 
formation of the flocks. For example, on one oc- 
casion in the Analalava Forest Zosterops, Nectar- 
inia souimanga, and N. notata flew in to within a 
few meters of the tape recorder seconds after the 
song of Calicalicus was played. 

Diet — The stomach of a bird taken 30 km 
NNW Fort-Dauphin (mnhn) contained "debris 
seulement animaux." Stomachs of our specimens 
contained spiders (Araneae), beetles (Curculioni- 
dae), Orthoptera, and Homoptera (Goodman & 
Parrillo, in press). 

Breeding — A male displayed to a female on 3 
November at 810 m in parcel 1 of the RNI 
d'Andohahela. The male followed the female 
closely for several minutes, fanning his tail wide- 
ly, spreading and drooping his wings, and calling. 
The male was in typical female plumage but with 
scattered black gular patches and thus was prob- 
ably molting from immature to adult plumage. 



80 



Another male observed the next day giving the 
same display was in full plumage. A male taken 
on 7 November in the Analalava Forest had testes 
8X6 mm (left, bilobed) and 4 X 3 mm (right). 

Weight— Male (2), 16.5, 18.0 g. 

Soft Part Colors — Bill: black with gray base 
and cutting edge; legs: gray; claws: black; iris: 
brown. 



Schetba rufa rufa 
Rufous Vanga 

The Rufous Vanga was recorded in humid for- 
ests from the Marovony Forest south through the 
Anosyenne and Vohimena mountains to the Man- 
antantely Forest between 50 and 850 m. In the 
humid forest of the RNI d'Andohahela (parcel 1) 
this species was only recorded at 440 and 810 m, 
where it was restricted to areas with large canopy 
trees on shallow slopes. Appert (1985) reported it 
from the Bemangidy Forest, and Bluntschli col- 
lected it near Eminiminy (amnh). The only record 
we have from littoral forest is one bird in October 
in the strand forest of Manafiafy. A specimen 
taken in 1756 was from near Fort-Dauphin, which 
is the type locality of this subspecies (Stresemann, 
1952; Rand, 1960). All material examined from 
southeastern Madagascar is referable to S. r. rufa. 

Diet — Stomachs of collected individuals con- 
tained spiders (Araneae: Salticidae), beetles (At- 
talabidae, Buprestidae, Cleridae, Elateridae, Scar- 
abaeidae, Staphylinidae), Hemiptera (Reduvi- 
idae), Homoptera, flies (Asilidae), ants (Formici- 
dae), and crickets (Gryllidae) (Goodman & 
Parrillo, in press). 

Breeding — On 27 October at 400 m in parcel 
1 of the RNI d'Andohahela, a pair was observed 
building a nest in a low fork of a tree 5 m from 
the ground. The nest was made of mosses and thin 
branches and was a deep cup shape. Both the male 
and the female brought nesting material. 

Adults collected in the Analalava Forest in early 
November and in the Marovony Forest in late Oc- 
tober and early November were in or approaching 
reproductive condition, including adult males with 
testes size ranging from 7X4 mm to 10 X 8 mm. 
In the Marosohy Forest in early December a pair 
of adults were observed feeding three fledglings. 

Weight— Female (2), 34.5, 40.5 g; male (3), 
38.0, 38.0, 40.0 g; combined (8), 36.8 ± 4.9 
(34.5-40.0) g. 

Soft Part Colors — Bill: cold grayish blue; 

FIELDIANA: ZOOLOGY 



legs: gray; claws: dark gray; iris: orangish red. 
Fledglings — Bill: dark gray with yellow mottling; 
legs: gray to cream colored; claws: gray; iris: 
brown. 



Vanga curvirostris curvirostris and Vanga 
curvirostris cetera 
Hook-billed Vanga 

The Hook-billed Vanga was recorded from the 
Marovony Forest south along the eastern coast 
and through the Anosyenne and Vohimena ranges 
to Petriky and west across the spiny forest to the 
Mandrare River, from sea level to 1200 m. It was 
the most widespread and locally common vanga 
observed at low elevations in southeastern Mad- 
agascar. It was rarely noted in groups of four or 
five individuals. In the RNI d'Andohahela (parcel 
1) this species was recorded at 440 and 1200 m, 
although it presumably occurred at low densities 
between these elevations. On 23 October, at 440 
m in parcel 1 of the RNI d'Andohahela, two birds 
were countersinging. Their calls provoked great 
agitation from nearby Phyllastrephus madagas- 
cariensis, Newtonia brunneicauda, Oxylabes 
madagascariensis, and Nectarinia souimanga, 
who approached the Vanga and called loudly. On 
another occasion in the same forest (13 Novem- 
ber, 1200 m), a silent V. curvirostris was being 
mobbed by a large group of passerines, including 
Neomixis, Newtonia spp., Leptopterus viridis, 
Cryptosylvicola randrianasoloi, and Dicrurus for- 
ficatus. 

This species is tolerant of habitat disturbance. 
For example, it nests in areas of spiny forest with 
extensive livestock damage and in monoculture 
stands of Pithecellobium duke. 

All specimens studied from humid and littoral 
forest sites in southeastern Madagascar are refer- 
able to V. c. curvirostris. The type locality of this 
subspecies has been restricted to Fort-Dauphin 
(Stresemann, 1952; Rand, 1960). Material from 
the spiny forest portion of the region collected in 
parcel 2 of the RNI d'Andohahela is referable to 
V. c. cetera (fmnh). 

Diet — Vanga primarily feeds on large insects 
and small vertebrates. At Manantantely this spe- 
cies has been observed taking young Brown 
Mouse Lemurs (Microcebus rufus) (Goodman et 
al., 1994b). In dry forests and to a lesser extent 
humid forests, chameleons make up a significant 
portion of this bird's diet. Including the tail, these 
chameleons may be almost as long as the bird 



itself. Chameleons are generally killed by holding 
them in the bill and whacking them against a 
branch. On several occasions Hook-billed Vangas 
wedged the large head of the dead chameleon into 
the fork of two branches, with the body of the 
chameleon hanging free below. On one occasion 
the chameleon was not dead when its head was 
wedged in the tree fork and was then dispatched 
by the bird by tugging at its body. The suspended 
carcass is then dismembered, piece by piece, by 
the vanga. The bird begins this process at the rear 
of the carcass, with the tail or hind limbs, often 
perching below the chameleon and reaching with 
the its heavy bill. Portions of the chameleon are 
fed to other individuals (presumably fledglings). 
The same butchering posts are regularly used by 
this species. Nestlings are also fed geckos, skinks, 
cockroaches, and a variety of other invertebrates. 

Cicadas often form a substantial portion of this 
vanga's diet, particularly during periods of emer- 
gence in November and December. The stomach 
contents of collected Vanga consisted of spiders 
(Araneae), centipedes (Geophilomorpha, Scolo- 
pendromorpha), flies (Asilidae), crickets (Grylli- 
dae), beetles (Chrysomelidae, Curculionidae, Ela- 
teridae, Scarabaeidae), cicadas (Cicadidae), true 
bugs (Psyllidae), Mantodea, ants (Formicidae), 
small amphibians, geckos (cf. Phelsuma), cha- 
meleons, and an unidentified passerine (Goodman 
& Parrillo, in press). 

Breeding — This species commences nest con- 
struction in August, and eggs hatch in late Sep- 
tember. During some years active nesting is noted 
during November and December; however, re- 
cords from these months may represent renesting 
by birds whose first clutch failed. A nestling was 
collected on 2 November at Amboasary-Sud 
(smf). The gonads of adults collected in mid-Sep- 
tember to late October ranged from small to large 
and in reproductive condition. An adult male 
taken on 17 October at Itapera had slightly en- 
larged gonads and a nonvascularized brood patch. 

The nests of Vanga are large, and nest place- 
ment is very stereotyped, which makes nests of 
Vanga among the easiest to find of any bird spe- 
cies in gallery and spiny forests. The open-cupped 
nests are lined with spider web (often Nephila) 
and usually lodged at the top of the trunk's bole 
at the point where the primary branches radiate 
from the main trunk. In several cases the same 
nest was reused over several consecutive breeding 
seasons, although it often was necessary for the 
vangas to repair the structure. In the RP de Ber- 
enty one nest was used for six consecutive sea- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



81 



sons. Preferred tree species for nest sites were 
Crateva excelsa, Celtis phillipensis, Pithecellob- 
iwn dulce, and, less frequently, Acacia rovumae, 
Ficus grevei, and Alluaudia procera (in spiny for- 
est). Nesting sites are in both closed-canopy and 
open-canopy forests. The clutch is usually three 
or four eggs. Both parents incubate and feed the 
young. Nests are heavily predated by Corvus al- 
bus, presumably because they are visible and ac- 
cessible to these crows. 

Weight — Vanga c. curvirostris female (2), 65, 
71 g; male (7), 71.6 ± 2.8 (68-75) g; combined 
(12), 71.0 ± 2.8 (65-75) g. V. c. cetera male (1), 
58.5 g. 

Soft Part Colors — Bill: black with whitish 
gray spot toward tip of maxilla; legs: dull bluish 
gray; claws: black and occasionally dark gray; 
iris: brown. 

Local Names — Fifiokala and tsilovanga (Ber- 
enty), vanga (Manafiafy, Manombo). 



Xenopirostris xenopirostris 
Lafresnaye's Vanga 

Lafresnaye's Vanga reaches its easternmost dis- 
tributional limit in southeastern Madagascar, 
where it is an uncommon inhabitant of the spiny 
forest portion from at least the Ankapoky Forest 
north to Hazofotsy and west to the Mandrare Riv- 
er, between near sea level and about 150 m. It 
was observed in both pristine and disturbed spiny 
forest. In the spiny forest adjacent to the Man- 
anara River, east of Hazofotsy, in the RNI 
d'Andohahela (parcel 2), probably no more than 
two or three pairs occurred along the 5 km of trail 
surveyed during the 1995 inventory of the region. 
A specimen was collected in late October at Ara- 
boasary-Sud (smf). 

This species is largely confined to spiny forest 
and the transitional zone between gallery and 
spiny forest. It regularly visits dead or dying trees 
with cracks or loose bark. It uses probing and lev- 
ering movements of its bill to extract insects, par- 
ticularly from various Euphorbia trees. 



Xenopirostris polleni 
Pollen's Vanga 

In southeastern Madagascar this species is 
largely confined to intact humid forest between 
440 and 1950 m. All of our records from the re- 
gion are from the Marosohy and Marosalohy for- 



ests and parcel 1 of the RNI d'Andohahela. Dur- 
ing the inventory of the latter forest parcel, this 
species was recorded between 810 and 1950 m 
but apparently was most common at 1 200 m. Mil- 
on collected a specimen 30 km NNW Fort-Dau- 
phin (mnhn). 

Between 27 November and 4 December, in 
transitional forest north of Trafonaomby, a mini- 
mum of two pairs were located. This species fre- 
quently foraged in low Philippia shrubs up to 100 
m from the edge of closed-canopy forest, where 
they were seen to catch and eat buprestid beetles. 
A male was observed on several occasions at the 
summit of Trafonaomby (1960 m), where vege- 
tation was low (about 1.5 m high) and dense and 
composed mostly of ericoid shrubs. This site is 
about 500 m higher than the previously reported 
elevational limit of this species (Langrand, 1990). 

On 29 October at 810 m in parcel 1 of the RNI 
d'Andohahela, a male X. polleni was seen mob- 
bing a Galidia elegans by swooping down to near 
the mammal's head and clacking its bill. While 
mobbing, the bird also gave a quiet "chuck- 
chuck-chuck" call. 

One juvenile was observed in close detail; it 
had a pale creamy orange underside (paler than 
an adult female) with a complete but not very 
extensive black hood (cf. Putnam, 1996). The bill 
was pale gray in the middle of both mandibles 
and pinkish along the culmen and cutting edges. 
The legs and feet were pale gray, paler than the 
adults', which were dark gray. The fleshy gape 
was obvious and pale orange, as was the narrow 
eye ring. 

On at least two occasions in the RNI 
d'Andohahela X. polleni was observed chasing 
Tylas eduardi. It is not clear why these species 
should display interspecific aggression; their for- 
aging methods are different (Yamagishi & Eguchi, 
1996; pers. obs.), although the plumage patterns 
of Tylas and female X. polleni are very similar. 

Diet — The stomach of a Milon specimen con- 
tained a large spider, caterpillar, and insect debris 
(mnhn). Like its congener X. xenopirostris of the 
spiny forest, X. polleni forages by searching for 
invertebrates concealed in woody substrates or 
among epiphytes. Oriolia bernieri, which, like X. 
polleni, is a vanga found in the eastern humid 
forest, also has very similar foraging behavior. 
Oriolia and X. polleni both climb up medium- 
sized or thick branches and strip off moss and 
loose bark, and both species characteristically 
perch on top of a horizontal branch and peer un- 



82 



FIELDIANA: ZOOLOGY 



derneath before tearing off some moss with the 
bill from the lower part of the branch. 

The distributions of Oriolia and X. polleni are 
imperfectly known, but they may be largely ex- 
clusive. Oriolia is found only in the northern por- 
tion of the eastern humid forest. The center of 
distribution of X. polleni appears to be in the 
southeast and east-central portions of the eastern 
humid forest, largely south of the range of Oriol- 
ia. The status of X. polleni within the range of 
Oriolia is not clear. There is a sight record of X. 
polleni from near Maroantsetra (Collar & Stuart, 
1985), where Oriolia is uncommon (Langrand, 
1990; pers. obs.). There is also a sight record of 
X. polleni from the RNI de Marojejy (Benson et 
al., 1977), although this species was not recorded 
there in a later survey (Safford & Duckworth, 
1990) or in the nearby RS d'Anjanaharibe-Sud 
(Hawkins et al., in press); Oriolia is known from 
both of these reserves. Perhaps the general pattern 
of largely exclusive distributions and the apparent 
rarity of X. polleni in areas occupied by Oriolia 
can be explained in part by their similar foraging 
techniques, which suggests that they might com- 
pete for food resources. 

Breeding — The testes of the Milon specimen 
collected on 26 May were 2 X 0.6 mm. The fol- 
lowing observations are from parcel 1 of the RNI 
d'Andohahela. On 15 November at 1200 m, a fe- 
male-plumaged bird was carrying food. On 9 No- 
vember at the same elevation, a probable imma- 
ture male (it had a blotchy pink breast and was 
speckled black on the central breast) was collect- 
ing leafy fibers, presumably for nesting material. 
On 29 November at 1 900 m, over a period of 40 
minutes, an adult male fed small insects five times 
to a juvenile. A female was in the vicinity but 
was not observed attending the juvenile during 
this feeding bout. The juvenile stayed in the same 
place for the whole period, calling continuously 
and quietly. 



Falculea palliata 
Sickle-billed Vanga 

In southeastern Madagascar the Sickle-billed 
Vanga is confined to the spiny forest, where it is 
often locally common in spiny and to a lesser ex- 
tent in gallery forest between 10 and 100 m. In 
gallery forest, this species prefers open areas with 
tall trees and little or no understory. It is more 
common in the Bealoka Forest than in the Malaza 



Forest. It was collected by Bluntschli near Am- 
boasary-Sud in late October (amnh, smf). 

Groups of up to 20 individuals were often ob- 
served as they moved noisily through the forest, 
sometimes with Leptopterus viridis. Falculea 
probes crevices in trunks and large branches and 
under bark with its long sickle-shaped bill (Ya- 
magishi & Eguchi, 1996; pers. obs.). On numer- 
ous occasions we observed Falculea perched or 
hanging upside down on the main trunks or ter- 
minal flower stalks of Alluaudia and, with side- 
sweeping motions of the bill, gleaning insects 
from the main trunk or inserting their bills into 
blossoms. 

Breeding — Falculea builds large nest struc- 
tures out of branches and twigs. The nests are 
often placed in Alluaudia or Adansonia trees and 
less often in Crateva excelsa trees. On 10 October 
at Ankapoky and on 15 November at Hazofotsy 
adult Falculea were observed refurbishing a nest 
that had been used during a previous breeding 
season. Nests are usually solitary structures, al- 
though on occasion several will be in the imme- 
diate vicinity of one another. For example, in No- 
vember 1984, five Falculea nests were in a tall 
isolated Crateva excelsa on the outskirts of Be- 
aloka. Two of the nests appeared to be abandoned, 
and three were in the process of being refur- 
bished. Several L. viridis entered the Falculea 
nests, shifted nesting material, and then flew off 
without damaging the nest or seemingly adding 
anything to the construction. The nesting and 
feeding relationships between Falculea and L vir- 
idis are in need of further investigation. 

Local Name — Voronzaza (Berenty). 



Leptopterus viridis viridis 
White-headed Vanga 

The White-headed Vanga occurs in humid for- 
est from Marovony south through the Anosyenne 
and Vohimena mountains at least to parcel 1 of 
the RNI d'Andohahela and then west into spiny 
forest to the Mandrare River, between 40 and 
1500 m. This species was not recorded in humid 
forest parcels (e.g., Bezavona and Manantantely) 
in the vicinity of Tolagnaro, although a specimen 
was collected in 1756 near Fort-Dauphin (Stre- 
semann, 1952). We also did not find this species 
in any littoral forest of southeastern Madagascar. 
In the 1960s or 1970s G. Randrianasolo recorded 
L. viridis at or near the Station Forestiere de Man- 
dena. The station, which is in littoral forest, is less 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



83 



than 2 km from the edge of the humid forest, and 
it is not clear if this species was observed in Man- 
dena proper. 

In the spiny forest area this species was rela- 
tively common in both intact and degraded spiny 
forest and in gallery forest. It occasionally was 
noted in relatively large groups, e.g., a flock of 
eight individuals on 5 September in the RP de 
Berenty (OL). Specimens collected near Amboas- 
ary-Sud (amnh, smf) in late October are referable 
to nominate viridis, the type locality of which has 
been designated as Tolagnaro (Stresemann, 1952; 
Rand, 1960). 

Frequently a few L. viridis were seen in flocks 
of Falculea, and these two species would forage 
together. Rather than using side-sweeping motions 
as Falculea, L. viridis pokes its bill into trunk crev- 
ices or into flowers. Tamarindus trees are regularly 
visited by these two species. L. viridis tends to 
probe into or lever away the tree's bark but is un- 
able to penetrate deeply into crevices and holes. In 
contrast, Falculea's sickle-shaped bill enables 
deeper penetration, but it is less adept at peeling 
bark from trunks. It is possible that Falculea ben- 
efits from the bark-loosening techniques of L. vir- 
idis. Indeed, Falculea may visit the exact foraging 
location of L. viridis as soon as the latter has de- 
parted. 

Diet — Stomach contents of collected birds in- 
cluded beetles (Chrysomelidae, Curculionidae), 
Homoptera, and ants (Formicidae) (Goodman & 
Parrillo, in press). 

Breeding — In parcel 1 of the RNI d' Andohahela 
on 27 October at 440 m, a male was seen gath- 
ering nest material, probably small sticks. A few 
days later at 810 m, a male was seen carrying 
food. In the RP de Berenty an isolated nest of this 
species, not in association with Falculea, was ac- 
tive between 16 and 21 November in a Neotina 
isoneura tree. 

Weight — Sex unknown (1), 50.5 g. 

Soft Part Colors — Bill: cold grayish blue at 
base merging to dull bluish gray at tip; mouth 
lining: black; legs: dark gray; iris: brown. 

Local Names — Vorompifkoka and bifeko (Be- 
aloka, Berenty). 



mena mountains to the Manantantely Forest and 
west to the Mandrare River, from 40 to 1000 m. 
This species was observed in both pristine and 
heavily disturbed humid and spiny forest and in 
villages and towns not far from the forest edge, 
e.g., Ranomafana-Sud. We did not record it in the 
littoral forests of Mandena, Manafiafy, or Petriky. 
However, G. Randrianasolo inventoried the birds 
near the Station Forestiere de Mandena in the 
1960s and 1970s and recorded L. chabert locally. 
The station is within 2 km of the humid forest, 
and it is not clear if this species was observed in 
Mandena proper. One individual was noted on 29 
May at Saihady, along this same coastal zone. 
There is a specimen collected in 1756 near Fort- 
Dauphin, which is the designated type locality of 
L. c. chabert (Stresemann, 1952; Rand, 1960). 

This species is rare in gallery forest, unless it 
is heavily disturbed. In dry forested areas, it is 
most frequently seen perched on and moving 
among Alluaudia procera trees. Flocks of up to 
12-14 individuals have been recorded in this re- 
gion. Specimens collected in the Ankapoky Forest 
are referable to L. c. schistocercus. Thus, the 
western slopes of the Anosyenne Mountains may 
form the divide between L. c. chabert and L. c. 
schistocercus. 

Diet — The stomachs of individuals collected 
near Ankapoky in mid-October contained beetle 
parts, seeds, and small fruits. 

Breedings — Individuals collected in mid-October 
at Ankapoky were in or approaching breeding con- 
dition. Adult males with large testes did not have 
brood patches. On 13 October in the Ankapoky 
Forest two adults were attending a small cup- 
shaped nest about 14 m off the ground attached to 
the main trunk of an Alluaudia tree. Three days 
earlier at a nearby locality five adult Chabert's Van- 
gas were observed flying around and alighting on 
a large stick nest about 10 m off the ground in a 
20-m-tall Alluaudia. The nest may well have orig- 
inally been constructed by Falculea. 

Weight — L. c. schistocercus female (1), 20.5 g; 
male (3), 18.0, 19.0, 19.0 g; combined (6), 19.0 ± 
0.8 (18.0-20.5) g. 

Soft Part Colors — Bill: bluish gray or whitish 
gray; legs and claws: black; iris: brown; orbital 
ring: mixture of brilliant sky blue and cobalt blue. 



Leptopterus chabert chabert and Leptopterus 
chabert schistocercus 
Chabert's Vanga 

Chabert's Vanga occurs from the Marovony 
Forest south through the Anosyenne and Vohi- 



Cyanolanius madagascarinus madagascari- 
nus 

Blue Vanga 

In southeastern Madagascar the Blue Vanga 
was relatively common at several intact and dis- 



84 



FIELDIANA: ZOOLOGY 



turbed humid forest sites. This species was fre- 
quently noted in clearings at the edge of the Be- 
zavona Forest. It has been reported from Beman- 
gidy (Appert, 1985). The Mandena Forest was the 
only littoral forest site in the area at which we 
recorded this species. In the spiny forest region it 
was relatively uncommon in spiny and gallery 
forest, e.g., the RP de Berenty. The elevational 
range of this species in southeastern Madagascar 
is from near sea level to 1500 m. The form oc- 
curring in the area is C. m. madagascarinus, the 
type locality of which has been designated as 
Fort-Dauphin (Stresemann, 1952; Rand, 1960). 

A multispecies flock encountered in parcel 1 of 
the RNI d'Andohahela at 1500 m on 26 Novem- 
ber contained five individuals, one of which was 
a juvenile that begged food from and was fed by 
an adult male. 

Diet — The stomach of a bird taken 30 km 
NNW Fort-Dauphin contained large insect re- 
mains (mnhn). 

Weight — Sex unknown (1), 22.5 g. 

Soft Part Colors — Bill: bright cold blue with 
black tip; legs: black; iris: dull blue. 



Hypositta corallirostris 
Nuthatch Vanga 

The Nuthatch Vanga was relatively uncommon 
in the region and confined to humid forest from 
the Marovony Forest south through the Anosyen- 
ne and Vohimena mountains to the Manantantely 
Forest. It was restricted to areas of intact humid 
forest between 75 and 650 m with large trees that 
were covered with mosses and epiphytes. In the 
RNI d'Andohahela (parcel 1) this species was 
only noted twice, at 440 m and 810 m. Appert 
(1985) recorded this species in the Bemangidy 
Forest. Salvan (1970) reported a specimen in the 
ORSTOM collection that was taken near Pic St. 
Louis but that cannot be traced. As of 1989 all of 
the appropriate habitat for this species on Pic St. 
1 -i mis had been destroyed. 

Peters (1996) described a new species, Hypos- 
itta perdita, on the basis of two Bluntschli spec- 
imens, both juveniles, taken near Eminiminy in 
September 1931. Peters was under the impression 
that H. corallirostris was not known from south- 
eastern Madagascar, although all of our sight re- 
cords from this region are of birds that appeared 
to be typical adult H corallirostris. Peters pro- 
posed, based on this interpretation of the foot 
structure of the two Bluntschli specimens, that H. 



perdita might not have the ability to grasp and 
climb tree branches and trunks. All of the Hypos- 
itta that we observed in southeastern Madagascar 
exhibited the bark-climbing behavior that is typ- 
ical of H. corallirostris. We know of no other ju- 
venile specimens of Hypositta, so the characters 
of perdita cannot be compared directly with cer- 
tain corallirostris of comparable age. Further 
study may show that the characters of H. perdita 
are those of juvenile H. corallirostris. 



Tylas eduardi eduardi 
Tylas Vanga 

In southeastern Madagascar the Tylas Vanga 
was largely confined to large tracts of humid for- 
ests between 350 and 1950 m. Our only records 
from the region were from the large forests in or 
adjacent to parcel 1 of the RNI d'Andohahela. 
During the 1995 inventory of the humid forest of 
this parcel, this species was observed at elevations 
from 440 to 1950 m, although it was more fre- 
quent at 440 and 810 m than at higher altitudes. 
A specimen was taken 30 km NNW Fort-Dauphin 
(mnhn). One individual was observed in heavily 
degraded forest at 940 m near Antseva. We have 
no records of this species from littoral forest. In 
the 1960s or 1970s, however, G. Randrianasolo 
recorded Tylas at the Station Forestiere de Man- 
dena. The station, which is in littoral forest, is less 
than 2 km from the edge of the humid forest, and 
it is not clear if this species was observed in Man- 
dena proper. Langrand (1990) reported Tylas from 
the general Tolagnaro area. 

There was considerable variation in plumage 
coloration of apparent adults. One presumed male 
observed in courtship feeding had almost com- 
pletely white underparts rather than the normal 
dark orange underparts. Another individual with 
a grayish white ventrum was seen at 810 m on 1 
November. Of a flock of four birds noted on 4 
November, three were very pale pinkish white un- 
derneath, and the coloration of the fourth could 
not be determined. One bird at 1500 m on 25 
November was completely white on the under- 
parts except for a single orange feather on the 
upper breast. 

An individual on 5 November at 810 m sunned 
itself by leaning its head over to one side, spread- 
ing the opposite wing and the tail, and remaining 
motionless in a patch of sunlight for several min- 
utes. A similar behavior has been observed in 
Schetba rufa. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



85 



Diet — One individual that caught a small ar- 
boreal cricket held the cricket in its foot as it 
leaned straight down and consumed the prey. 

Breeding — On 23 October at 440 m in the RNI 
d'Andohahela (parcel 1), a pair displayed to each 
other by wing-flicking (both wings were flicked 
downward and outward rapidly) and by courtship 
feeding. The specimen in mnhn taken on 28 May 
had testes 1.8 X 1.2 mm. 

Weight— Male (1), 43.5 g; combined (3), 43.5, 
46.0, 55.5 g; fledgling (1), 37.5 g. 

Soft Part Colors — Bill: black; legs: dull 
black or brownish black; claws: black; iris: brown 
and in one case greenish yellow. Fledgling — Bill: 
maxilla black with dull orange at base, and man- 
dible dull orange for most of its length with black 
tip; mouth lining and fleshy gape: bright yellowish 
orange; legs: dull gray; claws: black; iris: light 
brown with silver cast. 



Dicruridae 

Dicrurus forficatus forficatus 
Crested Drongo 

The Crested Drongo was one of the most wide- 
spread birds throughout southeastern Madagascar. 
This species was found in a broad variety of hab- 
itats from pristine to heavily disturbed littoral, hu- 
mid, and spiny forest, eucalyptus, pine, and sisal 
plantations, agricultural fields, river margins, and 
gardens in towns and villages between sea level 
and 1900 m. This bird was one of the few species 
that appear tolerant of human-induced habitat 
degradation. It was recorded at all of the QIT 
study sites and virtually every locality throughout 
southeastern Madagascar that we visited. It also 
has been reported from Bemangidy (Appert, 
1985). 

Diet — Stomachs of collected birds contained 
spiders (Araneae), beetles (Cerambycidae, Cur- 
culionidae, Elateridae, Scarabaeidae, Tenebrioni- 
dae), flies (Asilidae), Orthoptera, and Hymenop- 
tera (Goodman & Parrillo, in press). It is a highly 
versatile forager. It not only sallies into the air and 
to the ground from elevated perches but also will 
mob other species carrying captured prey. 

Breeding — Dicrurus nests typically in the open 
and constructs a small, shallow cup in which the 
adult barely fits. Adults are more often seen 
perched on top of rather than sitting in the nest 
cup. Several adults taken in mid-September to 
mid-October in the Manafiafy, Mandena, and 



Manantantely forests were in or approaching re- 
productive condition. In the Ankapoky Forest two 
adults were seen on 9 October attending a nest 5- 
6 m off the ground and placed in the fork between 
the main trunk and a horizontal branch. On 30 
October at 810 m in the RNI d'Andohahela (par- 
cel 1) an adult was on a nest about 8 m off the 
ground. In the dry areas of the region the breeding 
season extends from early October to mid-Feb- 
ruary. In the gallery forest along the Mandrare 
Dicrurus prefers to nest in the lower branches of 
large trees (Albizia, Acacia, Tamarindus, and 
Neotina). On 23 December two active nests were 
found in the RP de Berenty (OL), and on 26 De- 
cember in the Bezavona Forest two adults were 
seen feeding two free-flying fledglings. Males 
with greatly enlarged testes lacked brood patches. 

During the breeding season Dicrurus is excep- 
tionally territorial, and aggressive behavior to- 
ward conspecifics and other species of birds is 
relentless. It regularly is observed mobbing larger 
species such as Vanga, Eurystomus, Leptosomus, 
and virtually all birds of prey. Furthermore, Di- 
crurus is a convincing vocal mimic. At Bealoka 
it sometimes mimics Hypsipetes and Accipiter 
francesii. 

Weight— Female (1), 45 g; male (6), 49.2 ± 
2.8 (44-52) g; combined (15), 47.8 ± 4.7 (36- 
54.5) g. 

Soft Part Colors — Bill, mouth lining, legs, 
and claws: black; iris: dark reddish brown or 
crimson red. Fledglings — Mouth lining: yellow; 
iris: brown. 

Local Names — Lova (Berenty), railovy (Man- 
afiafy, Manombo, Marosohy). 



Corvidae 

Corvus albus 
Pied Crow 

In southeastern Madagascar the Pied Crow was 
typically found in areas along the eastern coastal 
plain and across the spiny forest region. The pres- 
ence of this species in the region was noted by 
Flacourt (1658) in the 17th century. It was almost 
exclusively found near human habitation, agricul- 
tural fields, and pastureland and occasionally 
along the seashore. The vast majority of records 
are between sea level and 50 m, although it is 
occasionally noted in degraded upland grassland 
areas such as near Eminiminy and Antseva. In the 
spiny forest region it does not occur in climax 



86 



FIELDIANA: ZOOLOGY 



forest but in open areas. It visits riverbanks on a 
daily basis, even during the hottest portion of the 
day. At dusk in areas surrounding spiny forest, a 
grand exodus of Pied Crows, sometimes number- 
ing several hundred, could be seen flying to 
roosts. This procession includes birds from the 
southern Mandrare basin and covers a period of 
about 1 hour. 

Corvus is often gregarious, particularly around 
concentrated food resources such as fish offal near 
fishermen's villages and discarded animal remains 
near slaughterhouses. It takes advantage of windy 
days to "hover" in the air, particularly along the 
coast and rivers. As with Dicrurus, Foudia mad- 
agascariensis, and Lonchura nana, this species 
may have benefited from human settlement of 
southeastern Madagascar. 

Diet — One individual collected at Mandena on 
1 September had insects, reptile bones, and sand 
in its stomach. In general it is a voracious nest 
thief in open areas. This species is probably re- 
sponsible for the failure of many nests of raptors 
in conspicuous locations, including Polyboroides, 
Buteo, and species such as Vanga curvirostris. In 
the RP de Berenty, Corvus takes advantage of or- 
thopteran invasions in meadow areas and also 
feeds extensively on the pericarps of the intro- 
duced Pithecellobium dulce tree. 

Breeding — The Mandena specimen was an 
adult female with an ovary 20 x 8 mm; the larg- 
est ovarian follicle was 3 mm. On 24 December 
an adult at Lac Anony was on a nest about 10 m 
up in a Casuarina tree at the edge of a barrier 
dune. In the RP de Berenty and Bealoka this spe- 
cies tends to place its nest toward the top of the 
tallest Acacia trees. 

Weight— Female (1), 580 g. 

Soft Part Colors — Bill, legs, and claws: 
black; iris: dark brown. 

Local Names — Goaka (Berenty), goaky (Man- 
ombo), kooky (Manafiafy). 



Sturnidae 

Hartlaubius auratus 
Madagascar Starling 

In southeastern Madagascar the Madagascar 
Starling was infrequently observed in a variety of 
habitats between 50 and 1200 m. It was most 
commonly noted in groups of two to four birds 
perched in the upper limbs of dead trees at the 
edge of humid forest, for example at Marovony, 



Analalava, Bemangidy, Marosohy, and Bezavona. 
On 27 December we observed this species near 
the western edge of parcel 1 of the RNI 
d'Andohahela, just below the Col d'Ambatomaniha 
in the ecotone between humid and dry forest. It 
also has been observed in littoral forest near Man- 
dena. Bluntschli collected this species near Emi- 
niminy in October (amnh, smf). There are speci- 
mens taken 7 km N Fort-Dauphin (mnhn) and in 
1756 near Fort-Dauphin (Stresemann, 1952). In 
the spiny forest region this species is uncommon. 

Diet — The stomach contents of a bird obtained 
near Mandena (mnhn) consisted of three seeds 
from a tree locally known as ambora. In the RNI 
d'Andohahela (parcel 1) a group of two to four 
Hartlaubius visited a site near our 810-m camp 
each morning, generally when it was sunny, to 
feed on the remains of an exploited beehive. 

Breeding — Specimens taken north of Tolagna- 
ro in late February (mnhn) had small reproductive 
organs. 



Acridotheres tristis 
Common Myna 

This introduced species is a relatively common 
human commensal throughout southeastern Mad- 
agascar in towns and villages and in other heavily 
modified habitats such as pastureland, agricultural 
fields, and gardens. Although the year of the ob- 
servations was not given, Sal van (1970) noted 
that sometime after 1945 this species was found 
30 km west of Tolagnaro and near Amboasary- 
Sud. In July 1982 it was found near Tolagnaro 
and in the RP de Berenty. We did not observe this 
species within the interior of any pristine forest, 
but it occurs at the edge of littoral, humid, gallery, 
and spiny forest. In heavily disturbed areas Acri- 
dotheres will penetrate forested habitats, often at 
considerable distances from human settlements. In 
the environs of Tolagnaro we observed flocks of 
up to 25 Common Mynas foraging on the ground 
near grazing cattle. 

This species is now widespread and has colo- 
nized the edge of natural forest areas and remote 
human settlements, e.g., Tsiombe, Ampanihy, and 
Ejeda. In the spiny forest this species nests in cav- 
ities of baobab trees, where it is more aggressive 
than native species utilizing the same holes and 
thus may be displacing them. This displacement 
may become a serious problem in areas such as 
the RP de Berenty, where numbers of Common 
Mynas have increased considerably in recent 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



87 



years, at least in part related to their utilization of 
food in open garbage pits. At this site it nests in 
the eaves of buildings. The subspecies introduced 
to the island is A. t. tristis. 

Diet — This species is largely omnivorous, and 
like Corvus albus, it is opportunistic, investigating 
anything and everything for palatability. 

Breeding — An adult male collected at Tolag- 
naro on 7 October had testes 8X6 mm (left) and 
6X5 mm (right). In the RP de Berenty this spe- 
cies has been found nesting during January and 
February. 

Weight— Male (1), 150 g. 

Soft Part Colors — Bill, legs, and claws: yel- 
low; iris: gray with white ring of spots; orbital 
ring: bright yellow. 

Local Names — Maritay (Manombo), ramaro 
(Manafiafy), rimaro (Berenty, Marosohy). 



Ploceidae 

Ploceus nelicourvi 
Nelicourvi Weaver 

The Nelicourvi Weaver is a relatively common 
bird of intact humid forests from the Marovony 
Forest south through the Anosyenne and Vohi- 
mena ranges to the Manantantely Forest between 
50 and 1950 m. In the RNI d'Andohahela (parcel 
1) this species was recorded in all zones between 
440 and 1950 m. We did not observe it in any 
littoral forest of southeastern Madagascar. How- 
ever, in the RS de Manombo, 140 km north of 
Manantenina, in an area where the littoral and hu- 
mid forests are separated only by a narrow cor- 
ridor of open ground, this species was found in 
both forest types. 

Diet — The stomach of a single collected bird 
contained spiders (Araneae), beetles (Curculioni- 
dae), and Homoptera (Goodman & Parrillo, in 
press). 

Breeding — Adult male and female specimens 
collected between late September and late Decem- 
ber often had enlarged gonads (e.g., testes up to 
10 X 6 mm). Fledglings were netted in the Ma- 
rovony Forest in late October and in the Bezavona 
Forest in late December. Males did not possess 
brood patches. This species generally nests soli- 
tarily, and the penduline nest is often attached to 
a tree limb and suspended over watercourses. This 
habit may be a adaptation to thwart snake preda- 
tion on nest contents. On 23 October, in parcel 1 
of the RNI d'Andohahela, a female-plumaged 



bird was singing near a nest. This bird may have 
been a young male rather than a female. In the 
same reserve at 1500 m on 25 November, another 
apparent immature male was carrying nest mate- 
rial. 

Weight— Female (6), 23.1 ± 1.1 (21.0-24.0) 
g; male (8), 24.6 ± 1.4 (23.0-26.5) g; combined 
(22), 23.9 ± 1.6 (20.5-26.5) g. 

Soft Part Colors — Bill: black; legs: brownish 
gray to black; claws: black and in a few individ- 
uals gray; iris: brown to reddish brown. Fledg- 
lings — Bill: black with random yellow patches. 

Local Names — Fody ala (Marosohy), fodisiay 
(Manombo). 



Ploceus sakalava minor 
Sakalava Weaver 

The Sakalava Weaver was a common inhabitant 
of the spiny forest of the region from near sea 
level to about 1 00 m. The easternmost records are 
from the east side of the Col de Ranopiso and the 
extreme western side of the RNI d'Andohahela 
(parcel 1), just above the village of Mahamavo, 
and to the west it occurs to the Mandrare River 
and beyond. All specimens examined from the re- 
gion are referable to P. s. minor. 

Diet — Stomach contents of collected individu- 
als invariably contained insect remains. In the An- 
kapoky Forest we observed this species picking 
at Alluaudia blossoms; they appeared to be eating 
the flowers. 

Breeding — Colonies of Sakalava Weavers were 
often placed in Adansonia za and Gyrocarpus 
americanus trees. The breeding regimen of this 
species appears to be irregular, presumably related 
to infrequent local rains rather than calendar sea- 
sons. In late August a colony was actively nest 
building in the middle of Anjapolo, 10 km north- 
west of Bealoka. On 15 November freshly 
hatched eggs were found below a colony near Ha- 
zofotsy. On 19 November birds collected from a 
colony near Bevilany consisted of an immature 
male with testes 6X4 mm and adult males and 
females in or approaching breeding condition. On 
23 December of the same year this colony was 
revisited, and remains of recently hatched eggs 
were found under the baobab tree. In mid-October 
numerous Sakalava Weavers in flocks were netted 
in the Ankapoky Forest. These flocks consisted of 
immatures, adult males with testes up to 10 X 8 
mm, and females with slightly enlarged ovaries. 
None of the males had a brood patch. One flock 



FIELDIANA: ZOOLOGY 



of + 30 birds, many of which carried nest mate- 
rial in their bills, was seen flying over the forest. 

According to local customs wealth and pros- 
perity are brought to a village if Sakalava Weav- 
ers choose to nest in a tree overhanging or beside 
a chief's house. For this reason this species is gen- 
erally left unmolested in villages. 

Weight— Female (6), 22.3 ± 2.6 (20.5-27.5) 
g; male (9), 25.1 ± 1.1 (23.5-27.0) g. 

Soft Part Colors — Bill: rather variable from 
mottled gray and black with light tip, light gray 
with cream-colored cutting edge to bluish gray; 
legs: grayish pink; claws: grayish pink; iris: 
brown, dull red, or dull orangish red; orbital ring: 
dull cream-colored or dull pinkish gray. 

Local Name — Fodibeotse (Berenty). 



Foudia madagascariensis 
Madagascar Red Fody 

The Madagascar Red Fody was one of the most 
common birds of the region from the Marovony 
Forest south along the coast and through the An- 
osyenne and Vohimena mountains to the southern 
coast and west across the spiny forest to the Man- 
drare River. It frequented a variety of habitats 
from the edge of littoral, spiny, and humid forests 
to heavily disturbed areas, including agricultural 
lands, pasturelands, thick secondary brush, and 
gardens, generally from sea level to about 1950 
m. In the spiny forest these birds are not common 
in intact gallery forest, and they usually occur in 
agricultural and degraded areas, often inhabiting 
the pediplain areas between forest, riverbank, and 
river. 

Immature birds, particularly males, and non- 
breeding adults congregated in flocks often num- 
bering up to 50 individuals and were observed in 
ripening grain fields and in gardens. Such a flock, 
netted in Tolagnaro on 4 October 1990, consisted 
of 1 3 individuals, all immatures with partially os- 
sified skulls; 12 were males and one was a female. 

Diet — Stomachs of collected individuals con- 
tained small insect parts and various types of 
seeds. 

Breeding — The breeding season of this species 
appears to be relatively protracted. For example, 
in the period from mid-September to late Novem- 
ber adults were taken in a variety of reproductive 
states, including active breeding, and during the 
same period subadults with partially ossified 
skulls were also obtained. A female taken on 28 
May, 30 km NNW Fort-Dauphin (mnhn), had an 



ovary 8.3 X 5.1 mm. In the dry parts of the region 
active nests have been found in May. 

Weight— Male (11), 17.2 ± 2.0 (13.5-19.5) g; 
combined (22), 16.9 ± 2.0 (13.0-19.5) g. 

Soft Part Colors — Bill: black and occasion- 
ally with dark gray mandible; legs and claws: 
pinkish brown, brown, or gray; iris: dark brown. 
Immatures — Bill: maxilla brown, and mandible 
brownish horn. 

Local Names — Folymena (Berenty), fody 
mena (Manafiafy, Manombo, Marosohy). 



Foudia omissa 
Forest Fody 

In southeastern Madagascar the Forest Fody 
was distinctly less common than the Madagascar 
Red Fody. Virtually all of our records of the For- 
est Fody come from within relatively large tracts 
of humid forest, e.g., Manantantely and Marosohy 
Forest and parcel 1 of the RNI d' Andohahela, be- 
tween 50 and 1900 m. Our only record from lit- 
toral forest was two birds netted at Mandena, one 
of which had an unossified skull (fmnh). At Man- 
antantely, this species was netted close to the for- 
est edge, about 500 m from where a pair of F. 
madagascariensis was captured. 

During the 1995 elevational transect of the RNI 
d' Andohahela (parcel 1), this species was found 
from 440 to 1900 m, although it was most fre- 
quently observed between 1200 and 1500 m. In 
this parcel it was often difficult to be certain of 
the identity of many individual Foudia; male F. 
madagascariensis may molt into breeding plum- 
age later in the year than F. omissa, and hence it 
may be difficult to distinguish between them. We 
have no clear evidence of hybridization between 
this species and F. madagascariensis, even in ar- 
eas where these two species are in close contact 
(see Benson et al., 1977). 

On 30 October, at 810 m in parcel 1 of the RNI 
d'Andohahela, in the late afternoon a dispersed 
group of 50-100 F. omissa was seen in and 
around a cluster of large Sloanea trees. There was 
much singing, calling, and flying around, and at 
least two separate groups of four or five males 
were seen displaying and calling together. This 
congregation may have been a roost grouping. 
Observations in this forest parcel and in other hu- 
mid forest sites of F. omissa flying over the forest 
canopy in the late afternoon suggest that at least 
during some portions of the year this species 
roosts communally. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



89 



I 



Diet — In parcel 1 of the RNI d'Andohahela 
birds were seen to feed on flowers of Strongylo- 
don, Bakerella, and Symphonia and on seeds of 
Sloanea. The stomachs of collected individuals 
contained some unidentified insects. 

Breeding — A pair of adults obtained on 1 Oc- 
tober from the same net and at the time consisted 
of a male with testes 8X6 mm and a female with 
a slightly enlarged oviduct, ovarian follicles up to 
4 mm in diameter, and no brood patch. 

Weight— Female (9), 18.6 ± 1.5 (17.0-21.0) 
g; male (9), 19.8 ± 1.6 (18.5-22.5) g; combined 
(18), 19.2 ± 1.6 (17.0-22.5) g. 

Soft Part Colors — Bill: maxilla black or 
dusky brown, and mandible black or yellowish 
brown with paler gonys; legs: pinkish gray or 
pinkish brown; iris: brown. 



Estrildidae 

Lonchura nana 

Madagascar Mannikin 

The Madagascar Mannikin was found from the 
Analalava Forest south along the coast and 
through the Anosyenne and Vohimena mountains 
to Petriky and west through the spiny forest to the 
Mandrare River. This species was occasionally 
noted in reed beds but generally was a character- 
istic bird of disturbed habitats, such as rice pad- 
dies, pasturelands, dry agricultural fields, and gar- 
dens from near sea level to 1150 m. It occasion- 
ally penetrated into disturbed areas within intact 
humid forests. For example, near the Col 
d' Ambatomaniha we found a flock of seven Mad- 
agascar Mannikins in an area of regenerating veg- 
etation associated with a natural landslide. Fur- 
ther, it is found in pockets of upland cleared areas, 
such as near Vohibaka and Antseva. It is distinctly 
less common in spiny forest than in more humid 
areas. Large flocks are occasionally noted, e.g., 
over 40 in the RP de Berenty in late September 
(DW). 

Diet — The diet is largely seeds. During May 
this species was observed feeding on the seeds of 
Phragmites. 

Breeding — Adults collected in mid-October 
and early November were not in breeding condi- 
tion. A pair was observed building a nest in To- 
lagnaro on 13 October, and another pair was ob- 
served mating at Bealoka on 4 December. Adults 
were noted feeding free-flying fledglings on 15 



October at Manafiafy and on 1 November near 
Antseva. 

Weight— Female (2), 8.5, 9.0 g. 

Soft Part Colors — Bill: maxilla black, and 
mandible silver gray; legs: grayish pink; claws: 
gray; iris: brown. 

Local Names — Fodimanta (Manombo), tsikin- 
itry (Manafiafy), tsimpiritry (Marosohy), tsipiritsy 
(Berenty). 



Analysis and Discussion 

General Overview of the Regional Avifauna 

Two hundred seventy-eight bird species have 
been recorded on Madagascar, 204 (73%) of 
which are known to breed on the island (Lan- 
grand, 1990; Langrand & Sinclair, 1994; Lan- 
grand & Appert, 1995; Goodman et al., 1996). 
Prior to the commencement of our field studies in 
southeastern Madagascar, 74 bird species were 
known to occur in this region. Currently, 189 spe- 
cies have been documented from southeastern 
Madagascar. A summary bird list for each of the 
study sites is presented in Table 6, along with in- 
formation on habitat association and geographic 
distribution. More detailed information for birds 
observed in the RNI d'Andohahela is presented 
by geographic (Table 7) and elevational (Table 8) 
distributions. 

Of the four littoral forest sites (Manafiafy, Itap- 
era, Mandena, Petriky), Manafiafy ranks the high- 
est in bird species richness (Table 6). The differ- 
ences in the number of species from the other 
three littoral forest sites are not pronounced. Like- 
wise, species richness in littoral forest does not 
differ markedly from that of adjacent humid forest 
(e.g., Bezavona compared with Mandena, and 
Manantantely compared with Petriky). 

The littoral forests tracts contained no bird spe- 
cies unique to that habitat. The near exception is 
Coua gigas, a species inhabiting the southwestern 
portion of the island and whose range extends 
through the littoral forests of southeastern Mada- 
gascar as far north as Manafiafy; this species does 
not occur in adjacent forests on lateritic soils. 

The humid forest sites (Marovony, Analalava, 
Marosohy, RNI d'Andohahela [parcel 1], Beza- 
vona, and Manantantely) had considerably higher 
species richness than the littoral forest sites. The 
relatively low species richness of the four littoral 
forest sites is even more pronounced considering 



90 



FIELDIANA: ZOOLOGY 






that all four sites contained aquatic habitats (sea- 
shore, coastal lagoons, etc.) that were not present 
at the six humid forests. The relative richness of 
the avifauna at 13 forested sites can best be as- 
sessed by restricting the analysis to birds that use 
forest habitats (Table 6). Such species can be di- 
vided into those that are forest dwellers (i.e., re- 
stricted to areas with closed forest canopy) and 
those that use mixed forest and open habitats. The 
four littoral forest sites contain fewer forest- 
dwelling bird species than do the humid forest 
sites. Below we comment on inventory conditions 
and sites and on interesting or unusual features of 
the avifauna of each surveyed forest. 

Mandena — The habitats surveyed at Mandena 
include littoral forest, swamp forest, heath (scrub), 
and marsh (estuary of Lac Ambavarano). At this 
site, the most comprehensive surveys were con- 
ducted within the littoral forests, concentrating 
mainly on the larger interior blocks where mist 
nets were erected and dawn censuses were con- 
ducted. The other littoral forest parcels were also 
visited. The forest parcel north of Pointe Evatra 
was highly degraded. 

Particularly noteworthy endemic species in- 
cluded Lophotibis cristata, Anas melleri, and 
Coua gigas, all three of which appeared to have 
been relatively common. Despite local hunting 
pressures on Lophotibis in the Mandena area, it 
was observed at least once per day at the site. 
Anas melleri utilized the aquatic habitats of the 
area. Relatively high densities of Coua gigas were 
noted in the forested areas. 

Petriky — Inventories at Petriky concentrated 
on littoral forests, although further surveys (by 
boat and on foot) were also made of estuarine 
aquatic habitats. Particularly interesting endemic 
species occurring at Petriky included Coua gigas 
and Ninox super ciliaris. The Coua was relatively 
common in the littoral forest. Petriky represents 
the easternmost known locality in southern Mad- 
agascar for Ninox. 

Manafiafy — Two distinct portions of Manafia- 
fy were intensively surveyed: a block of contin- 
uous littoral forest northwest of the village of 
! Manafiafy and a narrow strip of strand forest 
south of the village and across the Andohafasy 
River. The habitats in these two areas included 
littoral forest, swamp forest, heath (scrub), and 
marsh (waterways of the Andohafasy River). We 
I also visited a number of small remnant and dis- 
junct forest patches along the road leading from 
the Route Nationale 12a and a few kilometers 
! west of Manafiafy proper. 



Particularly noteworthy endemic species occur- 
ring in the area included Lophotibis cristata, Anas 
melleri, Coua gigas, and Asio madagascariensis. 
Relatively high densities of Coua gigas were not- 
ed, and it was restricted to the littoral forests. 
Manafiafy is the northernmost locality along the 
east coast of Madagascar at which this species has 
been recorded. 

Marosohy Forest — The two study sites within 
the Marosohy Forest were at different elevations 
(450 m and 750 m) along a trail leading from 
Antseva to Enakara that forms the northeastern 
boundary of parcel 1 of the RNI d'Andohahela. 
The two sites were in lowland forest. Mountain 
passes (up to nearly 1400 m) and lower agricul- 
tural areas at the edge of the reserve were also 
visited. 

Important endemic species occurring in the area 
were Lophotibis cristata, Mesitornis unicolor, 
Sarothrura insularis, Asio madagascariensis, 
Brachypteracias leptosomus, B. squamiger. Ate- 
lornis pittoides, Neodrepanis coruscans, Randia 
pseudozosterops, Newtonia fanovanae, and eight 
members of the Vangidae. 

Bezavona Forest — The Bezavona Forest 
drainage provides a portion of the potable water 
for Tolagnaro, and there is a pumping station at 
the base of the valley (see pp. 1 12-1 14 for further 
details). Endemic species occurring in the area of 
particular interest were Lophotibis cristata, Mes- 
itornis unicolor, Asio madagascariensis, and four 
members of the Vangidae. 

Analalava Forest — Interesting records of en- 
demic species occurring in this forest included 
Lophotibis cristata, Mesitornis unicolor, and three 
members of the Vangidae. 

Marovony Forest — The Marovony Forest 
contains an extensive lowland area on lateritic 
soils and is relatively undisturbed. This habitat is 
under intensive human pressure in southeastern 
Madagascar, and on a regional basis little of it 
remains (Green & Sussman, 1990). Noteworthy 
endemic species occurring in the Marovony For- 
est are Lophotibis cristata, Mesitornis unicolor, 
Caprimulgus enarratus, Philepitta castanea, and 
six members of the Vangidae. 

Manantantely Forest — The Manantantely 
Forest was surveyed between 75 and 200 m, al- 
though slopes of up to about 550 m were visited. 
Particularly interesting endemic species occurring 
in the area included Lophotibis cristata, Mesitor- 
nis unicolor, Brachypteracias squamiger, and 
four members of the Vangidae. 

The Manantantely Forest is particularly note- 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



91 



Table 6. Bird species recorded at each of the study sites in southeastern Madagascar, distributional status, and 
habitat preferences. 



















Site locations 3 














Sta- 
tus 1 


Habi- 
tat 2 


























Species 


ME 


PE 


SL 


MA 


NH 


AL MO MN 


IT 


AK 


ANIANII BE 


Phalacrocorax africanus 


N 


A 






















X 


X 


Ixobrychus minutus 


N 


A 


X 
























Nycticorax nycticorax 


N 


A 


X 


X 


X 










X 






X 


X 


Ardeola ralloides 


N 


A 




X 




X 














X 


X 


Ardeola idae 


* 


A 






















X 




Butorides striatus 


N 


A 




X 


X 










X 






X 


X 


Bubulcus ibis 


N 


O 


X 




X 


X 








X 






X 


X 


Egretta ardesiaca 


N 


A 






















X 




Egretta dimorpha 


(*) 


A 


X 




X 


X 








X 






X 


X 


Egretta alba 


N 


A 


X 


X 


X 


X 








X 


X 




X 


X 


Ardea purpurea 


N 


A 


X 


X 


X 


X 




X 




X 






X 


X 


Ardea cinerea 


N 


A 




X 


X 






X 


X 








X 


X 


Ardea humbloti 


(*) 


A 
























X 


Scopus umbretta 


N 


O 




X 




X 














X 




Lophotibis cristata 


* 


F 


X 




X 


X 


X 


X X 




X 




X 




X 


Phoenicopterus ruber 


N 


A 




X 






















Dendrocygna bicolor 


N 


A 






















X 


X 


Dendrocygna viduata 


N 


A 


X 


X 


X 


X 




X 




X 






X 


X 


Sarkidiornis melanotos 


N 


A 








X 














X 


X 


Anas melleri 


* 


A 


X 




X 




















Anas erythrorhyncha 


N 


A 


X 


X 


X 


X 








X 






X 


X 


Anas hottentota 


N 


A 




X 






















Aviceda madagascariensis 


* 


F 








X 


X 










X 


X 


X 


Machaeramphus alcinus 


N 


M 
























X 


Milvus migrans 


N 


O 


X 


X 


X 


X 


X 


X X 










X 


X 


Polyboroides radiatus 


* 


M 


X 


X 


X 


X 


X 


X X 


X 


X 


X 


X 


X 


X 


Accipiter madagascariensis 


* 


F 








X 




X 












X 


Accipiter francesii 


(*) 


F 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


Accipiter henstii 


* 


F 




















X 






Buteo brachypterus 


* 


M 


X 


X 


X 


X 


X 


X X 


X 


X 


X 


X 


X 


X 


Falco newtoni 


(*) 


O 




X 


X 


X 


X 


X X 






X 


X 


X 


X 


Falco zoniventris 


* 


F 


















X 




X 


X 


Falco eleonorae 


M 


O 




X 


X 
















X 


X 


Falco concolor 


M 


O 






X 
















X 


X 


Falco peregrinus 


N 


M 




X 




X 
















X 


Margaroperdix madagascarensis 


* 


O 


X 










X X 




X 




X 


X 


X 


Coturnix coturnix 


N 


O 
























X 


Numida meleagris 


N 


O 


X 


X 


X 


X 


X 


X X 




X 


X 




X 


X 


Mesitornis unicolor 


* 


F 








X 


X 


X X 


X 












Turnix nigricollis 


* 


M 


X 


X 


X 


X 


X 


X X 






X 


X 


X 


X 


Dryolimnas cuvieri 


(*) 


M 


X 




X 


X 


X 


X 




X 




X 


X 


X 


Canirallus kioloides 


* 


F 








X 


X 


X X 


X 


X 




X 






Sarothrura insularis 


* 


F 








X 












X 






Gallinula chloropus 


N 


A 


X 


X 


















X 




Porphyrio porphyrio 


N 


A 


X 




X 


















X 


Rostratula benghalensis 


N 


A 






X 


X 
















X 


Glareola ocularis 


* 


O 






X 


X 








X 










Charadrius hiaticula 


M 


A 






X 










X 










Charadrius thoracicus 


* 


A 
















X 










Charadrius pecuarius 


N 


A 




X 


X 










X 








X 


Charadrius tricollaris 


N 


A 












X 












X 


Charadrius leschenaultii 


M 


A 


X 
























Limosa lapponica 


M 


A 


X 
























Numenius phaeopus 


M 


A 






X 










X 








X 


Tringa nebularia 


M 


A 


X 


X 


















X 


X 


Actitis hypoleucos 


M 


A 




X 


X 


X 








X 






X 


X 


Arenaria interpres 


M 


A 






X 










X 










Gallinago macrodactyla 


* 


O 


X 










X 














Calidris alba 


M 


A 






X 










X 











92 



FIELDIANA: ZOOLOGY 



Table 6. Continued. 



















Site locations 3 














Sta- 
tus' 


Habi- 
tat 2 




























Species 


ME 


PE 


SL 


MA 


Ml 


A I. 


MO 


MN 


IT 


AK 


ANIAND BE 


Calidris ferruginea 


M 


A 


























X 


In/ it s dominicus 


N 


A 




X 


X 












X 








X 


Sterna caspia 


N 


A 


























X 


Sterna dougallii 


N 


A 






X 






















Sterna bergii 


N 


A 




X 
























Sterna bengalensis 


M 


A 






X 






















Pterocles personatus 


* 


O 




















X 




X 


X 


Streptopelia picturata 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Oena capensis 


N 


M 


X 


X 
















X 




X 


X 


Treron australis 


(*) 


F 


X 


X 


X 


X 


X 


X 


X 




X 




X 


X 


X 


Alectroenas madagascariensis 


* 


F 


X 




X 


X 


X 


X 


X 




X 




X 






Coracopsis vasa 


(*) 


F 








X 






X 






X 


X 


X 


X 


Coracopsis nigra 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Agapornis cana 


* 


M 


X 






X 






X 






X 




X 


X 


Cuculus rochii 


* 


F 








X 




X 


X 




X 


X 


X 


X 


X 


Coua gigas 


* 


F 


X 


X 


X 












X 


X 




X 


X 


Coua reynaudii 


* 


F 








X 


X 


X 


X 


X 






X 






Coua cursor 


* 


F 




















X 




X 


X 


Coua ruficeps 


* 


F 




















X 




X 


X 


Coua cristata 


* 


F 




X 
















X 




X 


X 


Coua caerulea 


* 


F 


X 




X 


X 


X 


X 


X 


X 


X 




X 






Centropus toulou 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Tyto alba 


N 


O 


X 
























X 


Otus rutilus 


(*) 


F 


X 






X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Ninox superciliaris 


* 


F 




X 
















X 




X 


X 


Asio madagascariensis 


* 


F 






X 


X 


X 




X 








X 




X 


Asio capensis 


N 


O 


























X 


Caprimulgus madagascariensis 


(*) 


O 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 




X 


X 


Caprimulgus enarratus 


* 


F 














X 








X 






Zoonavena grandidieri 


(*) 


S 








X 


X 


X 










X 


X 


X 


Cypsiurus parvus 


N 


S 


X 


X 


X 


X 




X 


X 




X 


X 


X 


X 




Apus melba 


N 


S 


X 


X 


X 


X 




X 


X 






X 


X 


X 




Apus barbatus 


N 


s 




X 




X 






X 






X 


X 




X 


Alcedo vintsioides 


(*) 


A 


X 


X 


X 


X 


X 


X 






X 




X 


X 


X 


Ispidina madagascariensis 


* 


F 


X 






X 


X 




X 


X 




X 


X 




X 


Merops superciliosus 


N 


O 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 


Eurystomus glaucurus 


N 


M 








X 


X 


X 


X 




X 


X 


X 


X 


X 


Brachypteracias leptosomus 


* 


F 








X 














X 






Brachypteracias squamiger 


* 


F 








X 








X 






X 






Atelornis pittoides 


* 


F 








X 














X 






Atelornis crossleyi 


* 


F 






















X 






Leptosomus discolor 


(*) 


F 




X 




X 


X 


X 


X 


X 






X 


X 


X 


Upupa epops 


N 


M 


X 


X 


X 












X 


X 




X 


X 


Philepitta castanea 


* 


F 








X 






X 








X 






Neodrepanis coruscans 


* 


F 








X 














X 






Neodrepanis hypoxantha 


* 


F 






















X 






Mirafra hova 


* 


O 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 


Riparia paludicola 


N 


S 






















X 






Phedina borbonica 


(*) 


S 








X 


X 








X 


X 


X 


X 


X 


Hirundo rustica 


M 


S 




X 
























Motacilla flaviventris 


* 


M 


X 




X 


X 


X 


X 


X 




X 




X 


X 


X 


Coracina cinerea 


(*) 


F 






X 


X 


X 


X 


X 


X 






X 


X 


X 


Phyllastrephus madagascariensis 


* 


F 








X 


X 


X 


X 


X 






X 






Phyllastrephus zosterops 


* 


F 








X 




X 




X 






X 






Phyllastrephus cinereiceps 


* 


F 






















X 






Hypsipetes madagascariensis 


N 


M 


X 


X 


\ 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Copsychus albospecularis 


* 


M 




X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Saxicola torquata 


N 


O 


X 


X 


X 


X 


X 


X 


X 








X 




X 


Pseudocossyphus sharpei 


* 


F 








X 














X 






Acrocephalus newtoni 


* 


A 
























X 


X 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



93 



Table 6. Continued. 



















Site locations 3 














Sta- 
tus' 


Habi- 
tat 2 




























Species 


ME 


PE 


SL 


MA 


NH 


AL 


MO 


MN 


IT 


AK ANIANII BE 


Nesillas lantzii 


* 


M 


? 


X 
















X 




? 


? 


Nesillas typica 


(*) 


M 






X 


X 


X 


X 


X 


X 


X 




X 






Thamnornis chloropetoides 


* 


M 
























X 


X 


Cisticola cherina 


* 


O 


X 


X 


X 


X 




X 


X 




X 


X 


X 


X 


X 


Dromaeocercus brunneus 


* 


F 






















X 






Randia pseudozosterops 


* 


F 








X 














X 






Cryptosylvicola randrianasoloi 


* 


F 






















X 






Newtonia amphichroa 


* 


F 








X 


X 


X 


X 








X 






Newtonia brunneicauda 


* 


F 




X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Newtonia archboldi 


* 


F 




















X 




X 


X 


Newtonia fanovanae 


* 


F 








X 














X 






Neomixis tenella 


* 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Neomixis viridis 


* 


F 








X 


X 












X 






Neomixis striatigula 


* 


F 








X 






X 






X 


X 


X 


X 


Hartertula flavoviridis 


* 


F 








X 


X 












X 






Pseudobias wardi 


* 


F 








X 














X 






Terpsiphone mutata 


(*) 


F 


X 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


Oxylabes madagascariensis 


* 


F 








X 


X 




X 


X 






X 






Crossleyia xanthophrys 


* 


F 






















X 






Mystacornis crossleyi 


* 


F 








X 


X 


X 


X 








X 






Nectarinia souimanga 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Nectarinia notata 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Zosterops maderaspatana 


(*) 


F 


X 






X 


X 


X 


X 


X 






X 


X 


X 


Calicalicus madagascariensis 


* 


F 








X 


X 


X 


X 


X 






X 




X 


Schetba rufa 


* 


F 






X 


X 




X 


X 








X 






Vanga curvirostris 


* 


F 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Xenopirostris xenopirostris 


* 


F 




















X 




X 


X 


Xenopirostris polleni 


* 


F 








X 














X 






Falculea palliata 


* 


F 




















X 




X 


X 


Leptopterus viridis 


* 


F 








X 






X 






X 


X 


X 


X 


Leptopterus chabert 


* 


M 








X 






X 


X 




X 


X 


X 


X 


Cyanolanius madagascarinus 


(*) 


M 


X 






X 


X 










X 


X 




X 


Hypositta corallirostris 


* 


F 








X 


X 




X 


X 






X 






Tylas eduardi 


* 


F 








X 














X 






Dicrurus forficatus 


(*) 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Corvus albus 


N 


O 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


X 


Hartlaubius auratus 


* 


M 


X 






X 


X 


X 


X 








X 






Acridotheres tristis 


I 


O 




X 


X 


X 




X 


X 




X 


X 




X 


x 


Ploceus nelicourvi 


* 


F 








X 


X 


X 


X 


X 






X 






Ploceus sakalava 


* 


F 




















X 




X 


X 


Foudia madagascariensis 


* 


M 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Foudia omissa 


* 


F 


X 






X 








X 






X 






Lonchura nana 


* 


O 




X 


X 


X 


X 


X 






X 






X 


X 


Total number of species at each site 






60 


59 


67 


100 


57 


58 


67 


37 


57 


52 


87 


85 


99 


Percentage of species endemic to Madagascar 


37 


22 


28 


53 


51 


48 


54 


60 


33 


50 


66 


38 


41 


Percentage of species restricted to region 




61 


45 


54 


77 


88 


81 


81 


97 


60 


79 


89 


61 


63 


Percentage of species nesting on Madagascar but 




























not endemic 






34 


45 


33 


20 


12 


17 


18 


3 


28 


19 


10 


30 


31 


Percentage of species that are migrants to area 


5 


5 


12 


1 














11 








4 


5 


Percentage of species introduced to 


Madagascar 





2 


2 


1 





2 


2 





2 


2 





1 


1 



1 * = endemic to Madagascar, (*) = endemic to region (Madagascar, Comoros, Mascarenes & Seychelles), M = 
migrant, N = nesting on Madagascar but not endemic to region, I = introduced. 

2 A = aquatic (including seashore), F = forest dwelling, O = open country, M = mixed forest and open country, 
S = aerial foraging. 

3 ME = Mandena Forest, PE = Petriky Forest, SL = Manafiafy Forest, MA = Marosohy Forest, NH = Bezavona 
or Nahampoana Forest, AL = Analalava Forest, MO = Marovony Forest, MN = Manantantely Forest, IT = Itapera, 
AK = Ankapoky Forest, ANI = RNI d'Andohahela (parcel 1), ANII = RNI d'Andohahela (parcel 2), BE - RP de 
Berenty. 



94 



FIELDIANA: ZOOLOGY 



Table 7. Avifaunal composition of study sites in the RNI d'Andohahela. 









Number (%) of species 












Endemic to 


Obligate 










Endemic to 


Malagasy 


forest 






Site 


Total 


Madagascar 


region 


species 


Migrants 


Introduced 


Andohahela (parcel 1) 














440 m 


60 


38 (63) 


56 (95) 


40(67) 








810 m 


65 


43 (66) 


61 (94) 


43(66) 








1200 m 


62 


42 (68) 


58 (94) 


42 (68) 








1500 m 


47 


30(64) 


44(94) 


36 (77) 








1875 m 


37 


23 (62) 


34 (92) 


24 (65) 








Overall 


87 


58 (67) 


78(90) 


52 (69) 








Andohahela (parcel 2) 


85 


35 (42) 


55 (65) 


23 (28) 


4(5) 


1(1) 



worthy for several reasons. It is positioned at the 
southern limit of the Vohimena Mountains and 
represents the southernmost large parcel of humid 
forest on the island. The Manantantely Forest is 
located at 24°59'S; thus it is south of the Tropic 
of Capricorn and, at least based on latitude, is out 
of the tropical zone. This forest is clearly tropical 
in structure and floristic components and is one 
of the southernmost "tropical" humid forests in 
the Old World. 

Itapera Forest — The habitats surveyed includ- 
ed the littoral forest, heath (scrub), and aquatic 
systems (associated with Lac Mananivo). The 
greatest effort was concentrated in the extant lit- 
toral forest, where mist nets were erected and 
dawn censuses were conducted. Particularly inter- 
esting endemic species found in the area included 
Lophotibis cristata, Charadrius thoracicus, and 
Coua gigas. Relatively high densities of C. gigas 
were noted in the area, and this species was re- 
stricted to the littoral forests. 

Ankapoky Forest — The Ankapoky Forest was 
the only site extensively studied during the QIT 
project in the spiny forest region. Of all the other 
QIT sites surveyed, Petriky is the closest to An- 
kapoky in both distance and floristic communities, 
although the flora of Petriky Forest is a transi- 
tional one between the eastern humid forest and 
spiny forest. Noteworthy endemic species occur- 
ring in the area include Falco zoniventris, four 
sympatric species of Coua, Newtonia archboldi, 
and six species of Vangidae. 

RNI d'Andohahela (Parcel 1) — The eastern 
slopes of the RNI d'Andohahela contain intact to 
pristine forest from approximately 400 to 1950 m. 
Our elevational transect of this forest included 
five sites along theses slopes centered at 440, 8 1 0, 
1200, 1500, and 1875 m. This region is rich in 
eastern humid forest species and marks the south- 



ern limit in the distribution of many species. 
Within the elevational transect, there was little 
proportional variation between zones in percent- 
age of the species endemic to Madagascar, species 
endemic to the region, or species that are forest 
dwelling (Tables 7, 8). The reserve has a rich avi- 
fauna, and particularly noteworthy endemic spe- 
cies included Asio madagascariensis, Caprimul- 
gus enarratus, all four humid forest species of 
Brachypteraciidae, both species of Neodrepanis, 
Randia pseudozosterops, the recently described 
Cryptosylvicola randrianasoloi, and nine species 
of Vangidae. 

RNI d'Andohahela (Parcel 2) — this parcel, 
composed largely of spiny forest region, contains 
an avifauna distinctly different from that of parcel 
1, although the two sites are separated by only a 
few kilometers. Interesting endemic species oc- 
curring in the area included Falco zoniventris, 
four sympatric species of Coua, Newtonia arch- 
boldi, and six species of Vangidae. 

Reserve Privee de Berenty — Although this 
privately owned reserve is small (about 250 ha), 
98 species of birds have been recorded there. The 
reserve is composed of spiny forest and gallery 
forest. The bird list is extensive because of the 
work conducted by MP over the course of several 
years and because numerous ecotourists and bird- 
watchers visit the site. This site, parcel 2 of the 
RNI d'Andohahela, and Ankapoky share many 
faunistic aspects. 



Relative Densities of Birds Based on Mist- 
netting 

Mist nets were used to assess relative densities 
of ground-dwelling and lower understory birds at 
most of the sites visited. There was considerable 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



95 



Table 8. Distribution of bird species in the Reserve Naturelle Integrate d'Andohahela. The elevational zones 
listed in the first five columns are along the 1995 transect between Eminiminy and Pic Trafonaomby. 

Parcel 1 



* 



Other 
Species 440 m 810 m 1200 m 1500 m 1875 m sites Parcel 2 

Phalacrocorax africanus * 

Nycticorax nycticorax * 

Ardeola ralloides * 

Ardeola idae * 

Bubulcus ibis * 

Butorides striatus * 

Egretta ardesiaca 
Egretta dimorpha 
Egretta alba 

Ardea purpurea * 

Ardea cinerea * 

Scopus umbretta * 

Lophotibis cristata * * 

Dendrocygna bicolor , * 

Dendrocygna viduata * 

Sarkidiornis melanotos * 

Anas erythrorhyncha * 

Aviceda madagascariensis * * 

Milvus migrans * 

Polyboroides radiatus * * 

Accipiter henstii * * 

Accipiter francesii * * * * * 

Buteo brachypterus ***** * 

Falco newtoni * * 

Falco zoniventris * 

Falco eleonorae * 

Falco concolor * 

Margaroperdix madagascarensis * * 

Numida meleagris * 

Turnix nigricollis * * 

Dryolimnas cuvieri * * 

Canirallus kioloides * * * * 

Sarothrura insularis * * * 

Gallinula chloropus * 

Tringa nebularia * 

Actitis hypoleucos * 

Pterocles personatus * 

Streptopelia picturata ***** * 

Oena capensis * 

Treron australis * * 

Alectroenas madagascariensis ***** 

Coracopsis vasa ***** * 

Coracopsis nigra ***** * 

Agapornis cana * 

Cuculus rochii ***** * 

Coua gigas * 

Coua reynaudii ***** 
Coua cursor * 

Coua ruficeps * 

Coua cristata * 

Coua caerulea ***** 

Centropus toulou * * * * * 

Otus rutilus * * * * * 

Ninox superciliaris * 

Asio madagascariensis * * * 

Caprimulgus madagascariensis * 

Caprimulgus enarratus * 

Zoonavena grandidieri * * * * * 

96 FIELDIANA: ZOOLOGY 



Table 8. Continued. 









Parcel 1 




















Other 




Species 


440m 


810 m 


1200 m 


1500 m 


1875 m 


sites 


Parcel 2 


Cypsiurus parvus 


* 


* 


* 








* 


Apus melba 


* 


* 


* 




* 




* 


Apus barbatus 


* 


* 


* 


* 


* 




* 


Alcedo vintsioides 


* 


* 










* 


Ispidina madagascariensis 


* 


* 


* 










Merops superciliosus 












* 


* 


Eurystomus glaucurus 














* 


Brachypteracias leptosomus 


* 


* 


* 










Brachypteracias squamiger 


* 














Atelornis pittoides 






* 










Atelornis crossleyi 




* 


* 




* 






Leptosomus discolor 


* 


* 


* 




* 




* 


Upupa epops 














* 


Philepitta castanea 


* 


* 


* 




• 






Neodrepanis coruscans 




* 


* 










Neodrepanis hypoxantha 






* 




* 






Mirafra hova 












* 


* 


Riparia paludicola 












* 




Phedina borbonica 




* 


* 




* 




* 


Motacilla flaviventris 


* 


* 


* 




* 






Coracina cinerea 


* 


* 


* 


* 


* 




* 


Phyllastrephus madagascariensis 


* 


* 


* 










Phyllastrephus zosterops 


* 


* 


* 










Phyllastrephus cinereiceps 




* 


* 


* 


* 






Hypsipetes madagascariensis 


* 


* 


* 


* 


* 




* 


Copsychus albospecularis 


* 


* 


* 




* 




* 


Saxicola torquata 












* 




Pseudocossyphus sharpei 




* 


* 


* 


* 






Acrocephalus newtoni 














* 


Nesillas lantzii 














? 


Nesillas typica 




* 


* 


* 


* 






Thamnornis chloropetoides 














* 


Cisticola cherina 












* 


* 


Dromaeocerus brunneus 






* 


* 


* 






Randia pseudozosterops 




* 




* 








Cryptosylvicola randrianasoloi 




* 


* 


* 


* 






Newtonia amphichroa 




* 


* 


* 


* 






Newtonia brunneicauda 




* 


* 


* 


* 




* 


Newtonia archiboldi 














* 


Newtonia fanovanae 
















Neomixis tenella 




* 


* 








* 


Neomixis viridis 




* 


* 


* 


* 






Neomixis striatigula 




* 


* 


* 






* 


Hartertula flavoviridis 




* 


* 


* 








Pseudobias wardi 




* 


* 


* 








Terpsiphone mutata 




* 


* 


* 






* 


Oxylabes madagascariensis 




* 


* 


* 


* 






Crossleyia xanthophrys 




* 


* 


* 


* 






Mystacornis crossleyi 




* 


* 


* 








Nectarinia souimanga 




* 


* 


* 


* 




* 


Nectarinia notata 




* 


* 








* 


Zosterops maderaspatana 




* 


* 


* 


* 




* 


Calicalicus madagascariensis 




* 


* 


* 








Schetba rufa 




* 












Vanga curvirostris 






* 








* 


Xenopirostris xenopirostris 














* 


Xenopirostris polleni 




* 


* 


* 


* 






Falculea palliata 














* 


Leptopterus viridis 


• 


* 


* 


* 






* 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



97 



Table 8. Continued. 









Parcel 1 




















Other 




Species 


440 m 


810 m 


1200 m 


1500 m 


1875 m 


sites 


Parcel 2 


Leptopterus chabert 


* 


* 


* 








* 


Cyanolanius madagascarinus 


* 


* 


* 


* 








Hypositta corallirostis 


* 














Tylas eduardi 


* 


* 


* 


* 


* 






Dicrurus forficatus 


* 


* 


* 


* 


* 




* 


Corvus albus 














* 


Hartlaubius auratus 


* 


* 


* 










Acridotheres tristis 














* 


Ploceus nelicourvi 


* 


* 


* 


* 


* 






Ploceus sakalava 














* 


Foudia madagascariensis 










* 




* 


Foudia omissa 


* 


* 


* 


* 


* 






Lonchura nana 














* 


Total number of species 


60 


65 


62 


47 


37 




85 



variation between sites in the number of species 
and individuals captured (Table 9). The results are 
analyzed by forest type. 

There were considerable differences between 
the number of species and individuals captured in 
the littoral forests of Petriky, Mandena, Manafia- 



Table 9. Summary of mist-netting results at study 
sites in southeastern Madagascar. Capture rate is defined 
as the number of individual birds captured per net-day. 







Num- 


Total 


Total 








ber 


num- 


num- 








of 


ber 


ber 








spe- 


of 


of 








cies 


12-m 


birds 


Cap- 




Forest 


cap- 


net- 


cap- 


ture 


Site 


type 


tured 


days 


tured 


rate 


Mandena 


Littoral 


12 


110 


24 


0.22 


Petriky 


Littoral 


3 


80 


3 


0.04 


Manafiafy 


Littoral 










North forest 




11 


240 


18 


0.08 


Strand 




13 


46.5 


42 


0.90 


Itapera 


Littoral 


2 


38.5 


3 


0.08 


Marosohy 


Humid 










425 m 




9 


50 


36 


0.72 


750 m 




20 


63 


79 


1.25 


Bezavona 


Humid 


11 


14 


24 


1.71 


Analalava 


Humid 


9 


31 


24 


0.77 


Marovony 


Humid 


22 


110 


82 


0.75 


Manantantely 


Humid 


14 


32 


37 


1.16 


Ankapoky 


Spiny 


17 


22 


96 


4.36 


Andohahela 


Humid 










440 m 




12 


50 


38 


0.76 


810 m 




21 


50 


61 


1.22 


1200 m 




17 


50 


48 


0.96 


1500 m 




16 


50 


61 


1.22 


1875 m 




9 


50 


41 


0.82 



fy, and Itapera (Table 10). At Manafiafy netting 
was conducted at two sites: strand forest and lit- 
toral forest. In the strand forest the capture rate 
was almost 10 times greater than in the littoral 
forest, although the number of species captured 
was approximately the same. At both sites the 
canopy height was approximately the same, and 
thus these differences may reflect absolute num- 
bers of understory birds at each site. In the forests 
of Petriky and Itapera few individuals or species 
were captured. Compared with the strand forest of 
Manafiafy, the species richness and relative den- 
sities at Itapera were remarkably low. Another 
comparison of these two areas comes from dawn 
censuses, during which 20 species were recorded 
at Manafiafy and 1 1 species were recorded at Itap- 
era (Table 11). On the basis of these results it 
appears that the relative density and species rich- 
ness in Itapera are distinctly lower than at Man- 
afiafy. These two sites are only a few kilometers 
from one another and are part of the same botan- 
ical formation. 

Considerably more species and four times as 
many individuals were captured at the spiny forest 
site of Ankapoky (Table 9). Ankapoky had the 
highest capture rate of any forest surveyed. The 
mist net capture rate of 4.4 individuals per net- 
day is inflated by the capture of 35 Ploceus sak- 
alava, a species locally restricted to this biome 
and moving in flocks at the time of the survey. 
Further, 18 Oena capensis were netted. These two 
species alone made up over half of the individuals 
captured. Even with these two species removed 
from the total net captures, about two individuals 



98 



FIELDIANA: ZOOLOGY 






Table 10. Mist-netting summary in the littoral forests of Petriky, Mandena, Manafiafy, and Itapera and spiny 
forest of Ankapoky. Entries are number of individuals captured/capture rate. 1 












Littoral (0-20 


m) 




Spiny 








Manafiafy 


Manafiafy 




Ankapoky 


Species 


Petriky 


Mandena 


strand 


north forest 


Itapera 


(70-100 m) 


Accipiter francesii 


1/0.01 


2/0.02 


1/0.02 


3/0.01 






Falco newtoni 












1/0.05 


Streptopelia picturata 




1/0.01 


2/0.04 


1/0.004 




2/0.09 


Oena capensis 












18/0.82 


Treron australis 






1/0.02 








Agapornis cana 












3/0.14 


Coua cristata 












1/0.05 


Otus rutilus 




1/0.01 










Caprimulgus madagascariensis 






5/0.11 






2/0.09 


Alcedo vintsioides 






1/0.02 


1/0.004 






Ispidina madagascariensis 




1/0.01 










Upupa epops 












2/0.09 


Hypsipetes madagascariensis 




1/0.01 


7/0.15 


1/0.004 




2/0.09 


Copsychus albospecularis 






2/0.04 


1/0.004 




8/0.36 


Cisticola cherina 












1/0.05 


Newton ia brunneicauda 






5/0.11 




1/0.03 


4/0.18 


Newtonia archboldi 












1/0.05 


Neomixis tenella 




1/0.01 


3/0.06 


2/0.008 






Neomixis striatigula 












1/0.05 


Terpsiphone mutata 


1/0.01 


1/0.01 


2/0.04 


3/0.01 






Nectarinia souimanga 


1/0.01 


9/0.08 


7/0.15 


4/0.02 


2/0.05 


5/0.23 


Nectarinia notata 




2/0.02 


5/0.11 


1/0.004 






Vanga curvirostris 




2/0.02 


1/0.02 


2/0.008 






Leptopterus chabert 












5/0.23 


Dicrurus forficatus 




1/0.01 




1/0.004 






Ploceus sakalava 












35/1.60 


Foudia omissa 




2/0.02 










Total net-days 


80 


110 


46.5 


240 


38.5 


22 


Total number captured 


3 


24 


42 


20 


3 


96 


Total number of species 


3 


12 


13 


11 


2 


17 



Capture rate is the number of individuals per species per net-day. 



were captured per net-day, a capture rate higher 
than at any other site in southeastern Madagascar 
(Table 9). The canopy of the Ankapoky Forest is 
distinctly lower than in any other Forest we stud- 
ied, and thus the height of the total flight corridor 
of birds was compressed. The high rate of net 
captures may not be entirely a measure of high 
density but rather may reflect our sampling of a 
greater portion of the forest cross section relative 
to canopy height. 

In lowland humid forests there was less varia- 
tion in the number of species and individuals cap- 
tured with mist nets (Tables 9, 12, 13). At the sites 
below 100 m (Bezavona, Analalava, Marovony, 
Manantantely), the number of birds captured per 
net-day ranged from 0.75 in the Maiovony Forest 
to 1 .7 1 in the Bezavona Forest. Although the Ma- 
rovony Forest had the lowest capture rate, more 
species were captured there than at the other three 



sites; this discrepancy is related to differences in 
the number of net-days at each site. 

Along the elevational transect of the humid for- 
est zone (parcel 1) of the RNI d'Andohahela stan- 
dardized mist-netting samples were obtained (Ta- 
ble 13). At each site (440, 810, 1200, 1500, and 
1875 m) 10 nets, 12 m long, were left in place 
for five consecutive days. Over the five zones 
there was little variation in capture rate, ranging 
from 0.76 at 440 m to 1.22 at both 810 and 1500 
m. The number of species netted in each zone was 
variable, with a peak at 810 m (21 species) and a 
low at 1875 m (nine species). This pattern is sim- 
ilar to that on a parallel transect conducted along 
the eastern humid forest slopes of the RNI 
d'Andringitra, where the highest netting success 
was in the 720-8 10-m zone and then success de- 
clined as a function of elevation (Goodman & Put- 
nam, 1996). However, in the RNI d'Andringitra 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



99 



Table 1 1 . Comparison of bird contact frequencies in the littoral forests of Manafiafy and Itapera based on dawn 
censuses. Relative density is measured as the number of individuals heard and observed per 100 m of trail. Total 
length of trails was not standardized. 







Manafiafy 






Itapera 






Relative 




Number 


Relative 




Number 


Species 


density 


(Range) 


of days' 


density 


(Range) 


of days 2 


Lophotibis cristata 


0.1 




1 


0.13 


(0.1-0.2) 


3 


Accipiter francesii 


0.4 




1 








Dryolimnas cuvieri 


0.15 


(0.1-0.2) 


2 








Streptopelia picturata 


0.16 


(0.1-0.3) 


3 


0.2 




1 


Treron australis 


0.1 




1 








Coracopsis nigra 


0.27 


(0.1-0.4) 


6 








Coua gigas 


0.20 


(0.1-0.4) 


4 


0.13 


(0.1-0.2) 


4 


Coua caerulea 


0.30 


(0.1-0.5) 


4 


0.1 




1 


Centropus toulou 


0.47 


(0.3-0.7) 


6 


0.2 


(0.2-0.2) 


2 


Merops superciliosus 


0.15 


(0.1-0.2) 


2 


0.1 




1 


Coracina cinerea 


0.20 


(0.2-0.2) 


2 








Hypsipetes madagascariensis 


0.19 


(0.1-0.4) 


6 


0.2 




1 


Copsychus albospecularis 


0.10 


(0.1-0.1) 


4 


0.1 




1 


Newtonia brunneicauda 


0.25 


(0.1-0.6) 


6 


0.3 


(0.1-0.6) 


3 


Neomixis tenella 


0.69 


(0.2-1.3) 


7 








Terpsiphone mutata 


0.13 


(0.1-0.2) 


4 








Nectarinia souimanga 


1.16 


(0.7-1.7) 


7 


1.3 


(1.1-1.4) 


4 


Nectarinia notata 


0.1 




1 








Vanga curvirostris 


0.28 


(0.1-0.6) 


5 


0.4 


(0.3-0.6) 


3 


Dicrurus forficatus 


0.15 


(0.1-0.2) 


2 









1 Number of mornings out of 7 the species was recorded. 

2 Number of mornings out of 4 the species was recorded. 



there was no sample in the 400-m zone and the 
710-m site included partially open and disturbed 
forest with some human hunting pressure. The 
number of species captured in the RNI 
d'Andohahela and RNI d'Andringitra at 810-m 
sites were 21 and 13, respectively, at 1200 m and 
1210 m were 17 and 13, respectively, and at 1500 
m and 1625 m were 16 and 15, respectively. 

Birds also were netted in the Marosohy Forest 
at 425 and 750 m, which is along the northern 
edge of the RNI d'Andohahela (parcel 1) and is 
part of the same large tract of forest. The nets at 
the 425 -m site in the Marosohy Forest captured 
10 species and 36 individuals during 50 net-days, 
whereas at 440 m in the RNI d'Andohahela 12 
species and 38 individuals were captured during 
60 net-days. At the 750-m site in the Marosohy 
Forest 20 species and 79 individuals were cap- 
tured in 63 net-days, whereas at 810 m in the RNI 
d'Andohahela 21 species and 61 individuals were 
captured in 50 net-days. Thus, there were broad 
similarities between these two sites in relative 
numbers of individuals and species captured in 
nets. 

When sites with netting data are combined by 
forest type, capture rates (weighted average, 



100 



range) are: littoral and strand forest, 0.17, 0.04- 
0.9; spiny forest, 4.36; and humid forest, 1.10, 
0.72-1.71. Thus, the spiny forest had the highest 
capture rate of any forest type, and more under- 
story birds were captured in the taller humid for- 
ests than in the littoral forests. 



Elevational Distribution of Birds 

The species richness of forest birds declined 
with elevation in parcel 1 of the RNI 
d'Andohahela (Table 8), as expected from bird 
studies in other tropical areas (Terborgh, 1971, 
1977; Prigogine, 1980; Beehler, 1982; Goodman 
et al., 1995). Comparative data from other eleva- 
tional transects in Madagascar, at the RS 
d' Anjanaharibe-Sud and the RNI d'Andringitra, at 
both of which the elevational samples started at 
about 800 m, also show a similar pattern of a 
decline in species number above this elevation. 
The 440-m sample in the RNI d'Andohahela had 
five fewer species than the 800-m sample. Be- 
cause this difference was relatively small and be- 
cause forests below 400 m are largely destroyed 
(which may influence species richness at this site), 

FIELDIANA: ZOOLOGY 






Table 12. Mist-netting summary in the lowland humid forests of Marosohy, Bezavona, Analalava, Marovony, 
and Manantantely. Entries are number of individuals captured/capture rate. 1 















Manan- 




Marosohy 


Marosohv 


Bezavona 


Analalava 


Marovony 


tantely 


Species 


(425 m)* 


(750 m)" 


(75 m) 


(40 m) 


(50 m) 


(-100 m) 


Accipiter francesii 




1/0.015 


1/0.07 




1/0.01 




Streptopelia picturata 




1/0.015 


1/0.07 






1/0.03 


Otus rutilus 




2/0.03 








2/0.06 


Caprimulgus madagascariensis 










3/0.03 




Caprimulgus enarratus 










1/0.01 




Alcedo vintsioides 




1/0.015 










Ispidina madagascariensis 


1/0.02 


4/0.06 


1/0.07 




1/0.01 


1/0.03 


Merops superciliosus 










1/0.01 




Brachypteracias leptosomus 




1/0.015 










Philepitta caslanea 


9/0.18 


11/0.17 






1/0.01 




Neodrepanis coruscans 




1/0.015 










Phyllastrephus madagascariensis 


4/0.08 


11/0.17 




8/0.26 


15/0.14 


7/0.22 


Phyllastrephus zosterops 


3/0.06 


3/0.05 




1/0.03 






Hypsipetes madagascariensis 


2/0.04 


5/0.08 


6/0.43 




6/0.05 


7/0.22 


Copsychus albospecularis 


5/0.10 


9/0.14 


2/0.14 


2/0.06 


15/0.14 


5/0.16 


Nesillas typica 










2/0.02 




Newtonia amphichroa 










1/0.01 




Newtonia brunneicauda 




1/0.015 




1/0.03 


1/0.01 


5/0.16 


Newtonia fanovanae 


1/0.02 












Hartertuia flavoviridis 


2/0.04 












Terpsiphone mutata 


7/0.14 


10/0.16 




6/0.19 


9/0.08 




Oxylabes madagascariensis 










6/0.05 




Mystacornis crossleyi 




1/0.015 


1/0.07 




2/0.02 




Nectarinia souimanga 




2/0.03 


1/0.07 


1/0.03 


2/0.02 


1/0.03 


Nectarinia notata 








3/0.10 


3/0.03 




Zosterops maderaspatana 


2/0.04 


6/0.1 


1/0.07 


2/0.06 


4/0.04 


3/0.09 


Calicalicus madagascariensis 












1/0.03 


Schetba rufa 








3/0.10 


1/0.01 




Dicrurus forficatus 




1/0.015 


1/0.07 




1/0.01 


2/0.06 


Ploceus nelicourvi 




8/0.13 


7/0.14 




5/0.01 


1/0.03 


Foudia madagascariensis 






2/0.14 




1/0.01 


2/0.06 


Foudia omissa 












2/0.06 


Total net-days 


50 


63 


14 


31 


110 


32 


Total number captured 


36 


79 


24 


31 


82 


37 


Total number of species 


10 


19 


11 


11 


22 


14 



1 Capture rate is the number of individuals per species per net-day. 



we do not consider this evidence of a mid-eleva- 
tion peak in species richness (sensu Janzen et al., 
1976; McCoy, 1990; Olson, 1994). 

The only site in Madagascar that has been stud- 
ied in a manner comparable to that of the RNI 
d'Andohahela is the RNI de Zahamena, with data 
from 500 to 1500 m. At this site, the 500-m sam- 
ple was by far the most species rich (Hawkins, 
unpubl.). In contrast, the 440-m sample at RNI 
d'Andohahela had five fewer species than the 
sample at 800 m. Differences in the bird com- 
munity between the RNI d'Andohahela and RNI 
de Zahamena provide an additional explanation 
for the slightly lower species richness of lowland 
forest at Andohahela. The only bird species sig- 
nificantly more abundant in the RNI d'Andohahela 



than in the RNI de Zahamena were Crossleyia 
xanthophrys and Xenopirostris polleni. Both spe- 
cies were most common in the RNI d'Andohahela 
above 800 m. In contrast, five species were re- 
corded in the RNI de Zahamena that were not 
found in the RNI d'Andohahela (Eutriorchis as- 
tur, Coua serriana, Phyllastrephus tenebrosus, 
Oriolia bernieri, and Euryceros prevostii). All 
were recorded in the RNI de Zahamena only at 
500 m, and elsewhere in their ranges they are un- 
known or very rare above 1000 m (Thorstrom & 
Watson, 1994; Hawkins et al., in press). Thus, the 
major difference in the bird community between 
parcel 1 of the RNI d'Andohahela and sites to the 
north is in the lowland species. Most of these spe- 
cies are not known south of the region south of 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



101 



Table 13. Mist-netting summary of birds captured along an elevational gradient in the RNI d'Andohahela (parcel 
1). Recaptures are not included. Entries are number of individuals captured/capture rate. 1 









Elevation (m) 






Species 


440 


810 


1200 


1500 


1875 


Streptopelia picturata 








2/0.04 




Otus rutilus 




1/0.02 








Alcedo vintsioides 




3/0.06 








Ispidina madagascariensis 


2/0.04 


5/0.10 


1/0.02 






Brachypteracias squamiger 


1/0.02 










Atelornis pittoides 






2/0.04 






Philepitta castanea 


4/0.08 


5/0.10 


7/0.14 


10/0.20 


2/0.04 


Neodrepanis coruscans 




3/0.06 


1/0.02 






Neodrepanis hypoxantha 








4/0.08 


4/0.08 


Motacilla flaviventris 




3/0.06 








Phyllastrephus madagascariensis 


8/0.16 


1/0.02 


3/0.06 






Phyllastrephus zosterops 


4/0.08 


3/0.06 


7/0.14 






Phyllastrephus cinereiceps 






1/0.02 


4/0.08 




Hypsipetes madagascariensis 


2/0.04 


4/0.08 


1/0.02 


2/0.04 




Copsychus albospecularis 


6/0.12 


5/0.10 








Pseudocossyphus sharpei 




2/0.04 




1/0.02 


4/0.08 


Nesillas typica 




1/0.02 


1/0.02 


8/0.16 


8/0.16 


Newtonia amphichroa 






4/0.08 


8/0.16 


1/0.02 


Newtonia brunneicauda 










1/0.02 


Hartertula flavoviridis 








2/0.04 




Terpsiphone mutata 


6/0.12 


7/0.14 


4/0.08 


3/0.06 




Oxylabes madagascariensis 




1/0.02 


1/0.02 


4/0.04 




Crossleyia xanthophrys 






3/0.06 


1/0.02 




Mystacornis crossleyi 


1/0.02 










Nectarinia souimanga 


1/0.02 






1/0.02 


12/0.24 


Zosterops maderaspatana 


1/0.02 


9/0.018 




1/0.02 


6/0.12 


Leptopterus viridis 




1/0.02 








Cyanolanius madagascariensis 




1/0.02 








Tylas eduardi 




1/0.02 


3/0.06 






Dicrurus forficatus 




3/0.06 


1/0.02 






Ploceus nelicourvi 


2/0.04 


1/0.02 


2/0.04 


3/0.06 




Foudia omissa 




1/0.02 


6/0.12 


7/0.14 


3/0.06 


Total net-days 


50 


50 


50 


50 


50 


Total number captured 


38 


61 


48 


61 


41 


Total number of species 


12 


21 


17 


16 


9 



Capture rate is the number of individuals per species per net-day, not including recaptures. 



the PN de Ranomafana, and their absence from 
the RNI d'Andohahela reflects a decrease in hu- 
mid forest bird species richness with increasing 
latitude. 



Utilization of Sisal Plantations by Forest 
Birds 

During forest surveys in the gallery forests of 
Malaza and Bealoka along the Mandrare River 
within the spiny forest zone in 1984 and 1985 it 
was noticed that many birds made use of sisal 
(Agave rigida) during flowering and fruiting. 
These two forests are now small remnants of the 
former natural habitat and are surrounded by 



thousands of hectares of sisal plantations. In its 
sterile form sisal provides little sustenance or cov- 
er for most birds (Fig. 15), but when flowering it 
becomes a sought-after resource, both for food 
(directly by providing flowers, pollen, and nectar 
and indirectly by attracting insects) and as favored 
perching sites from which hunting, displaying, 
mating, and sunning may take place. 

Agave inflorescenses are tall and over a period 
of a few months may grow to over 6 m in height. 
An Agave plant may take 20-30 years to accu- 
mulate enough carbohydrates to support one sea- 
son of flowering (Crawford, 1989). In a commer- 
cial setting, plantations are usually large enough 
and with sufficient different generations of plants 
to provide a substantial and predictable food re- 



102 



FIELDIANA: ZOOLOGY 




Fig. 15. Young sisal plantation near Amboasary-Sud. A relatively high diversity of birds use such areas, but only 
when the plants are in flower. (Photograph by T. S. Schulenberg.) 



source from year to year even though each plant 
flowers only once in its lifetime. However, the 
usefulness of sisal to forest birds as a resource is 
greater when close to forest. 

Sisal plantations border the western boundaries 
of the Mala/a Forest and the Bealoka Forest, 
which is 7 km farther north along the Mandrare 
River. Monthly data were collected by MP on 
birds utilizing sisal at both sites between May 
1984 and April 1985. Perimeter transects were 
walked one day per month for 1 year. All birds 
seen or heard, as well as height off ground, type 
of activity, distance from observer, and substrate 
preference (i.e., perched on inflorescence or 
among leaves), were recorded. 

At the Malaza Forest the 1-km perimeter trail 
bordering the sisal was used for the transect. 
The survey in this forest started in June 1984. 
Transects were walked in the morning and eve- 
ning until September, after which transects were 
walked only in the morning. A total of 20.5 
hours, covering 15 km, was spent surveying the 
Malaza sisal plantation. At Bealoka, a sisal 
plantation bordered the forest along the com- 



plete length of the 1.5-km trail. The survey here 
commenced in May 1984 and followed the same 
procedure during the period of the study. Totals 
of 25.5 km of trail and 33 hours of observations 
were accrued at this site. 

Twenty-nine species of birds were recorded in 
the sisal plantations during the survey, and a fur- 
ther 10 species were observed in the sisal at other 
times. Of these 39 species, 25 (63.9%) are con- 
sidered forest-dwelling species (Fig. 16). Using 
the results from both forests, feeding was one of 
the most common activities. Thirty-three percent 
of birds observed during the transects were feed- 
ing from sisal flower heads either directly or in- 
directly (Table 14). At Malaza however, most 
birds were observed perched on the sisal and vo- 
calizing. Feeding we defined as birds engaged in 
the process of consuming at the moment of the 
observation. Birds undertaking "food searches" 
were not considered to be feeding. 

The height, stability, and food offered by the 
inflorescences was an important enticement for 
birds to visit sisal. Collectively (at Malaza and 
Bealoka) only 7.8% of birds were observed in the 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



103 



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Bealoka 



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co co 

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to 

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5 


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CO 


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CI 




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CD 

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co 

3 


CJ 


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3 


o 

.CJ 


CO 

a 

v. 


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<£ •£ S 



•2 £ .« co 

c co a *. 



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ill 



a co s 



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2. ■ 



o 

rjv .CO Co -O 

co 



m co -5 2 



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t: Co 
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CJ 



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I 



Fig. 16. Total number of individuals per species recorded in sisal plantation along survey transects at the edge 
of the Malaza and Bealoka forests. Species recorded utilizing sisal but not observed during transect surveys included 
Buteo brachypterus, Falco peregrinus, Coracopsis vasa, Coua gigas, Upupa epops, Newtonia brunneicauda, Neomixis 
striatigula, Terpsiphone mutata, Leptopterus viridis, and Ploceus sakalava. 



Table 14. Sisal use by birds' adjacent to the Malaza and Bealoka forests. 





Total 


Total 


Perched 


Perched 






Average 




number 


diver- 


on 


on 


Perched 


Perched 


height 


Months 


of birds 


sity 


inflorescence 


foliage 


vocalizing 


feeding 


(m) 


1984 
















May 


— /82 


— ■ no 


— /78 


— /3 


— /35 


— /33 


— /4.5 


June 


13/62 


3/8 


6/62 


7/0 


11/14 


0/22 


3.4/4.7 


July 


19/29 


5/10 


19/29 


0/0 


7/6 


7/10 


5.1/4.7 


August 


60/40 


9/9 


57/40 


3/0 


28/4 


26/30 


5.8/4.6 


September 


17/20 


6/7 


16/20 


1/0 


5/3 


9/8 


4.3/4.1 


October 


29/17 


10/5 


26/17 


2/0 


11/1 


6/0 


3.7/4.8 


November 


2/11 


2/6 


1/11 


1/0 


0/2 


0/3 


4.5/5.2 


December 


17/12 


6/7 


13/8 


4/4 


6/3 


0/0 


3.7/3.5 


1985 
















January 


2/10 


1/4 


2/6 


0/4 


2/3 


0/0 


6.0/3.8 


February 


2/3 


2/2 


0/3 


2/0 


2/0 


0/0 


1.8/3.6 


March 


4/2 


2/2 


0/2 


4/0 


0/0 


0/0 


0.4/3.4 


April 


4/6 


2/2 


3/6 


1/0 


5/1 


0/0 


4.2/5.0 


Total 


169/294 


18/25 


143/282 


25/11 


77/72 


48/106 


3.9/4.3 


Total by percentage 






84.6/95.9 


14.8/3.7 


45.6/24.5 


28.4/36.1 




Combined results 


463 


29 2 


425 


36 


149 


154 


4.1 


Combined results by 
















percentage 






91.8 


7.8 


32.2 


33.3 





1 Presented as number at Malaza/number at Bealoka. 

2 An additional 10 species were observed in sisal outside of count periods. 



104 



FIELDIANA: ZOOLOGY 






foliage, and at Bealoka only 3.79c were observed 
in the foliage (Table 14). Although eight species 
of birds were observed perched among the sisal 
foliage, 50% of observations were represented by 
only two species: Centropus toulou and Cisticola 
cherina. The former is a common but rather se- 
cretive skulking bird that benefits from the cover 
the sisal leaves provide. The latter is an inhabitant 
of the grass layer and roadside verges; it was seen 
only in sisal foliage and never up high on the 
flower stalk. 

Direct comparisons of sisal utilization along the 
two transects are difficult. At Malaza along the 
shorter transect the sisal was only one or two 
plants wide along the entire survey route and di- 
rectly abutted the forest periphery. At Bealoka ex- 
tensive sisal fields of hundreds of hectares border 
the forest and the two habitats are separated by a 
10-m-wide road, which creates a barrier between 
forest and plantation that some forest birds may 
not cross. The rewards are greater for the birds of 
Bealoka than for those of Malaza because of the 
more extensive sisal fields. Nevertheless, birds 
perched in sisal at Bealoka presumably were more 
exposed to predation, which may partly explain 
why 80% of all observations in sisal at this site 
were less than 30 m from the forest edge. 

In 1984 sisal was flowering between May and 
October. Although the results presented in Table 
14 suggest a decline in feeding after October, this 
is just as much a consequence of demand by local 
people, who use the stems for fence poles, as it 
is a period of flower degeneration. Following the 
removal of the inflorescence stalks there was a 
distinct change in bird activity. The frequency of 
birds declined substantially, and at Malaza more 
birds were seen in foliage than in inflorescences. 
During the last 4 and 5 months 50% of all birds 
were recorded or seen over 30 m from the forest, 
and these birds were mostly "open country" spe- 
cies, i.e., Falco concolor, Centropus toulou, Nec- 
tarinia souimanga, Hypsipetes rnadagascariensis, 
Cisticola cherina, Dicrurus forficatus, Acridoth- 
eres tristis, and Foudia rnadagascariensis. The 
most accessible flower stalks, those generally 
closest to trails and roadways, were felled, and the 
removal of this resource caused an instant reduc- 
tion in the number of birds along the transect. 

Apart from perching and feeding in flower 
heads and using the foliage for cover, birds oc- 
casionally exploited plant sap from the sisal's 
growing stem. In July and August 1984, perhaps 
the period of maximum flowering that year and 
also during the middle of the dry season, Lemur 



catta at Malaza would climb halfway up the flow- 
er spikes and chew holes into the sides of the stem 
to feed on the rising sap. These holes were 
worked on daily until hollows within the stem 
held considerable quantities of liquid sap. This re- 
source was exploited by Hypsipetes, Neomixis te- 
nella, Nectarinia souimanga, and Zosterops. At 
times the sunbirds were seen drinking from the 
same holes in sisal stems as beetles, wasps, and 
flies. 

In conclusion, sisal's usefulness to birds is 
maximized when it grows adjacent to forest. A 
noticeable decline in bird species richness in sisal 
occurs as a function of distance from the forest 
edge. Sisal is principally utilized by birds during 
inflorescence growth, when it provides food and 
elevated perches. Sisal in its sterile state is of little 
appeal to forest birds and provides only a tem- 
porary habitat for some ground-dwelling birds 
and birds of open country. 



Faunistics and Biogeography 

Within southeastern Madagascar there is a re- 
markable change in the climate and flora across 
the rain shadow caused by the Anosyenne Moun- 
tains. Here we examine the faunal affinities of the 
forest-dwelling bird fauna within southeastern 
Madagascar in comparison with the bird com- 
munities of humid and dry forest sites elsewhere 
on the island. We define forest-dwelling species 
as those that occur in forest but are not necessarily 
forest dependent. Opposite extremes of this defi- 
nition include the ground-rollers (Brachypteraci- 
idae), which are completely forest dependent, and 
generalists, such as Hypsipetes rnadagascariensis 
and Nectarinia spp., which occur in a wide variety 
of habitats, including forest. 

We have chosen sites for this analysis that are 
ornithologically well known and represent a wide 
range of different forest types (Table 15). To as- 
sess the relationships of the regional and extra- 
limital avifauna, we calculated two different sim- 
ilarity indices for the distribution of breeding 
birds within forest habitat: 



Simpson's Index = — 



Jaccard's Index = 



C 



/V, + N 2 - C 
where /V, - the number of species at site 1 (the 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



105 



Table 15. Distribution of resident forest-dwelling birds 1 at well-known sites in the eastern humid forest, spiny 
bush, and transitional forest. Key to sites: AnI = RNI d'Andohahela (parcel 1), And = RNI d'Andringitra (restricted 
to humid forest), Ran = PN de Ranomafana, Anl = RS d'Analamazaotra, Anj = RS d'Anjanaharibe-Sud, Mas = 
proposed PN de Masoala, Mda = Montagne d'Ambre, Anil = RNI d'Andohahela (parcel 2), Ber = RP de Berenty, 
Bez = RS de Beza Mahafaly, Zom = proposed PN de Zombitse. Entries in brackets denote that the local subspecies 
is inferred. 



Species 



Humid forest 



Subarid thorn 
scrub 



Tran- 
si- 
tional 



Anl And Ran Anl Anj Mas Mda Anil Bez Zom 

(400- (720- (750- (900- (860- (0- (900- (80- Ber (100- (750- 

1900 1625 1100 1000 1950 1224 1450 150 (15- 200 900 

m) m) m) m) m) m) m) m) 30 m) m) m) 



Lophotibis c. cristata 


X 


X 


X 


X 


X 


X 


X 










Lophotibis c. urschi 


















[x] 




[x] 


Aviceda madagascariensis 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Machaeramphus alcinus 












X 






X 






Eutriorchis astur 








X 


X 


X 












Polyboroides radiatus 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


Accipiter madagascariensis 


X 


X 


X 


X 


X 


X 


X 




X 


X 




Accipiter f. francesii 


X 


X 


X 


X 


X 


X 


X 




X 


X 


X 


Accipiter henstii 


X 


X 


X 


X 


X 


X 


X 








X 


Buteo brachypterus 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Falco n. newtoni 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Falco zoniventris 




X 




X 


X 


X 




X 


X 


X 


X 


Falco peregrinus radama 


X 


X 






X 








X 






Mesitornis unicolor 


X 


X 


X 


X 


X 


X 












Turnix nigricollis 


X 


X 








X 




X 


X 


X 


X 


Dryolimnas c. cuvieri 


X 




X 


X 


X 


X 


X 


X 


X 


X 


X 


Canirallus k. kioloides 


X 


X 


X 


X 


X 


X 


X 










Sarothrura insularis 


X 


X 


X 


X 


X 


X 


X 










Pterocles personatus 
















X 


X 


X 


X 


Streptopelia p. picturata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Oena capensis aliena 














X 


X 


X 


X 


X 


Treron a. australis 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Alectroenas madagascariensis 


X 


X 


X 


X 


X 


X 


X 










Coracopsis v. vasa 


X 


X 


X 


X 




X 


X 










Coracopsis v. drouhardi 
















[x] 


[X] 


[X] 


[X] 


Coracopsis n. nigra 


X 


X 


X 


X 


X 


X 


X 










Coracopsis n. libs 
















[x] 


[x] 


[x] 


[x] 


Agapornis c. cana 


X 


X 








X 












Agapornis c. ablactanea 
















X 


X 


X 


X 


Cuculus audeberti 




X 




X 
















Cuculus rochii 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Coua gigas 
















X 


X 


X 


X 


Coua serriana 








X 


X 


X 












Coua reynaudii 


X 


X 


X 


X 


X 


X 












Coua cursor 
















X 


X 


X 




Coua ruficeps olivaceiceps 
















X 


X 


X 


X 


Coua c. cristata 






X 


X 


X 


X 


[X] 










Coua c. pyropyga 
















X 


X 


X 


X 


Coua caerulea 


X 


X 


X 


X 


X 


X 












Centropus t. toulou 


X 


X 


X 


X 


X 


X 




X 


X 


X 


X 


Tyto soumagnei 








X 


X 


X 


X 










Otus r. rutilus 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Ninox superciliaris 












X 


X 


X 


X 


X 


X 


Asio madagascariensis 


X 


X 


X 


X 




X 


X 




X 


X 


X 


Caprimulgus enarratus 


X 




X 


X 


X 


X 


X 










Zoonavena g. grandidieri 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Ispidina m. madagascariensis 


X 


X 


X 


X 


X 


X 


X 




X 






Ispidina m. diluta 






















X 


Eurystomus g. glaucurus 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Brachypteracias leptosomus 


X 


X 


X 


X 


X 


X 













106 



FIELDIANA: ZOOLOGY 



Table 15. Continued. 



Humid forest 



Subarid thorn 
scrub 



Tran- 
si- 
tional 



Species 



AnI And Ran Anl Anj Mas Mda Anil Bez Zom 

(400- (720- (750- (900- (860- (0- (900- (80- Ber (100- (750- 

1900 1625 1100 1000 1950 1224 1450 150 (15- 200 900 

m) m) m) m) m) m) m) m) 30 m) m) m) 



Brachypteracias squamiger 


X 


X 






X 


X 












Atelornis pittoides 


X 


X 


X 


X 




X 


X 










Atelornis crossleyi 


X 


X 


X 


X 


X 


X 












Leptosomus d. discolor 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Upupa epops marginata 














X 


X 


X 


X 


X 


Philepitta castanea 


X 


X 


X 


X 


X 


X 












Neodrepanis coruscans 


X 


X 


X 


X 


X 


X 












Neodrepanis hypoxantha 


X 


X 




X 


X 














Coracina c. cinerea 


X 


X 


X 


X 


X 


X 


X 










Coracina c. pallida 
















X 


X 


X 


[X] 


Phyllastrephus m. madagascariensis 


X 


X 


X 


X 


X 


X 


X 










Phyllastrephus m. inceleber 






















X 


Phyllastrephus z, zosterops 


X 


X 


X 


[X] 
















Phyllastrephus z- fulvescens 














X 










Phyllastrephus z. maroantsetrae 












X 












Phyllastrephus z- andapae 










X 














Phyllastrephus apperti 






















X 


Phyllastrephus tenehrosus 








X 




X 












Phyllastrephus cinereiceps 


X 


X 


X 




X 


X 












Hypsipetes m. madagascariensis 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Copsychus a. albospecularis 










X 


X 


[x] 










Copsychus a. pica 
















X 


X 


X 


X 


Copsychus a. inexpectatus 


X 


X 


X 


X 


X 














Pseudocossyphus s. sharpei 


X 


X 


X 


X 


X 


X 












Pseudocossyphus s. erythronotus 














X 










Pseudocossyphus bensoni 






















X 


Nesillas t. typica 


X 


X 


X 


X 


X 


X 


X 










Nesillas lantzii 
















X 


X 


X 


X 


Thamnornis chloropetoides 
















X 


X 


X 


X 


Dromaeocercus brunneus 


X 


X 


X 


X 


X 














Dromaeocercus seebohmi 




X 


X 


















Randia pseudozosterops 


X 


X 


X 


X 


X 














Cryptosylvicola randrianasoloi 


X 


X 


X 


X 


X 














Newtonia amphichroa 


X 


X 


X 


X 


X 


X 


X 










Newtonia b. brunneicauda 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Newtonia archboldi 
















X 


X 


X 




Newtonia fanovanae 


X 








X 


X 












Neomixis t. tenella 










[X] 


X 


X 










Neomixis t. debilis 




















[x] 


[x] 


Neomixis t. orientalis 


X 


X 


[X] 


[x] 








[x] 


[x] 






Neomixis viridis subsp. 


X 


X 


X 


X 


X 


X 












Neomixis striatigula sclateri 


X 


X 


X 


X 


X 


X 












Neomixis s. pallidior 
















X 


X 


X 


X 


Hartertula flavoviridis 


X 


X 


X 


X 


X 














Pseudobias wardi 


X 


X 


X 


X 


X 


X 












Terpsiphone m. mutata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Oxylabes madagascariensis 


X 


X 


X 


X 


X 


X 


X 










Crossleyia xanthophrys 


X 


X 


X 


X 


X 














Mystacomis crossleyi 


X 


X 


X 


X 


X 


X 












Nectarinia s. souimanga 


X 


X 


X 


X 


X 


X 


X 










Nectarinia s. apolis 
















X 


X 


X 


X 


Nectarinia n. notata 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Zosterops m. maderaspatana 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Calicalicus madagascariensis 


X 


X 


X 


X 


X 


X 


X 




X 




X 


Schetba r. rufa 


X 


X 


X 


X 


X 


X 













GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



107 



Table 15. Continued. 



Humid forest 



Subarid thorn 
scrub 



Tran- 
si- 
tional 



Species 



AnI And Ran Anl Anj Mas Mda Anil Bez Zom 

(400- (720- (750- (900- (860- (0- (900- (80- Ber (100- (750- 

1900 1625 1100 1000 1950 1224 1450 150 (15- 200 900 

m) in » in ) m I 111 1 in ) m) in I 30 in I m) in i 



Schetba r. occidentalis 






















X 


Vanga c. curvirostris 


X 


X 


X 


X 


X 


X 


X 










Vanga c. cetera 
















X 


X 


X 


X 


Xenopirostris xenopirostris 
















X 


X 


X 


X 


Xenopirostris polleni 


X 


X 


X 


X 




X 












Falculea palliata 
















X 


X 


X 


X 


Leptopterus v. viridis 


X 


X 


X 


X 


X 


X 












Leptopterus v. annae 
















[x] 


[x] 


[x] 


tx] 


Leptopterus c. chabert 


X 


X 


X 


X 


X 


X 


X 










Leptopterus c. schistocercus 
















X 


[X] 


[x] 


[x] 


Cyanolanius m. madagascarinus 


X 


X 


X 


X 


X 


X 


X 




X 




X 


Oriolia bernieri 










X 


X 












Euryceros prevostii 








X 


X 


X 












Hypositta corallirostris 


X 


X 




X 


X 


X 












Tylas e. eduardi 


X 


X 


X 


X 


X 


X 


X 










Dicrurus f. forficatus 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Hartlaubius auratus 


X 


X 


X 


X 


X 


X 


X 








X 


Ploceus nelicourvi 


X 


X 


X 


X 


X 


X 


X 










Ploceus sakalava minor 
















X 


X 


X 


X 


Foudia madagascariensis 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Foudia omissa 


X 


X 


X 


X 


X 


X 













1 Sources of information: Anl = herein, includes Marosohy Forest; And = Goodman and Putnam (1996); Ran = 
Nicoll and Langrand (1989), Goodman et al. (1996), Goodman (unpubl.); Anl = Nicoll and Langrand (1989), Lan- 
grand and Sinclair (1994), Goodman et al. (1996); Anj = Nicoll and Langrand (1989), Halleux and Goodman (1994), 
Hawkins et al. (in press); Mas = Nicoll and Langrand (1989), Watson and Strzalkowska (1992), Thorstrom and 
Watson (1994), Langrand and Sinclair (1994); MDA = Langrand (1995), Andrianarimisa (1994), Andrianarimisa and 
Goodman (unpubl.), B. Freed (pers. comm.), S. M. Goodman (unpubl.); Anil = herein, Nicoll and Langrand (1989); 
Ber = herein, Nicoll and Langrand (1989); Bez = Nicoll and Langrand (1989), Ratsirarson (1996), Goodman (un- 
publ.); Zom = Goodman et al. (1994a). 



smaller fauna), N 2 = the number of species at site 
2, and C = the number of species common to both 
sites. The coefficients from these indices were 
used in a cluster algorithm (PHYLIP, using the 
Fitch-Margoliash method with contemporary tips 
written by J. Felsenstein). 

Further, to demonstrate two different aspects of 
the biogeographic relationships and species turn- 
over across the island, and specifically across the 
pluviometric fault, we calculated the two indices 
using two different species concepts: the biolog- 
ical species (Mayr, 1942, 1992) and the phylo- 
genetic species (McKitrick & Zink, 1988) con- 
cepts. In the context of these analyses, distinct 
geographic forms (subspecies) of a taxon are 
combined as a single taxonomic unit of a biolog- 
ical species and separated as different taxonomic 
units of a phylogenetic species. We have chosen 
the latter approach to emphasize presumed genetic 



isolation between distinct populations of the east- 
ern humid forest and those of the dry areas to the 
west. The coefficients of the analyses for the bi- 
ological species are presented in Table 16 and 
those of the phylogenetic species are presented in 
Table 17. 

Affinities of the Regional Avifauna — All 
four analyses show that the avifaunas of the hu- 
mid forests and the dry forests are distinctly dif- 
ferent from one another and that sites of compa- 
rable forest type form close clusters (Fig. 17), re- 
gardless of geographic distance. On a regional ba- 
sis the two habitat types within the RNI 
d'Andohahela (which in southeastern Madagascar 
exemplify differences between the wet and dry 
forests), although only separated by about 5 km 
of ground distance (our study sites within these 
parcels are about 30 km apart), have the most bio- 
geographically distinct avifaunas of any sites cho- 



108 



FIELDIANA: ZOOLOGY 






Table 16. Faunal similarity indices using a biological species concept of resident forest-dwelling birds of several 
well-known sites. Above the diagonal is Jaccard's Index and below the diagonal is Simpson's Index. 1 Locality ab- 
breviations follow Table 15. 





Anl 


And 


Ran 


Anl 


Anj 


Mas 


Mda 


Anil 


Ber 


Bez 


Zom 


AnI 





0.93 


0.89 


0.84 


0.88 


0.79 


0.58 


0.36 


0.43 


0.38 


0.36 


And 


0.% 


— 


0.86 


0.86 


0.78 


0.77 


0.54 


0.36 


0.43 


0.39 


0.45 


Ran 


0.97 


0.96 


— 


0.87 


0.82 


0.77 


0.64 


0.37 


0.39 


0.40 


0.48 


Anl 


0.92 


0.94 


0.97 


— 


0.86 


0.83 


0.60 


0.34 


0.43 


0.37 


0.44 


Anj 


0.95 


0.88 


0.95 


0.93 


— 


0.83 


0.55 


0.32 


0.36 


0.34 


0.41 


Mas 


0.90 


0.88 


0.92 


0.92 


0.91 


— 


0.61 


0.39 


0.45 


0.41 


0.48 


Mda 


0.91 


0.87 


0.92 


0.94 


0.89 


0.96 


— 


0.45 


0.49 


0.47 


0.53 


Anil 


0.70 


0.70 


0.66 


0.68 


0.66 


0.77 


0.66 


— 


0.84 


0.90 


0.76 


Ber 


0.71 


0.71 


0.64 


0.73 


0.64 


0.75 


0.68 


1.00 


— 


0.89 


0.79 


Bez 


0.74 


0.72 


0.70 


0.80 


0.66 


0.75 


0.64 


0.98 


1.00 


— 


0.78 


Zom 


0.63 


0.74 


0.74 


0.74 


0.70 


0.79 


0.72 


0.96 


0.89 


0.94 


— 



1 See text (p. 105) for definitions of indices. 



sen for this analysis. The bird fauna of the eastern 
humid forest, a region extending nearly 1,200 km 
between parcel 1 of the RNI d'Andohahela and 
the RS d'Anjanaharibe-Sud, is more homoge- 
neous than that of the few kilometers separating 
parcel 1 and parcel 2 of the RNI d'Andohahela. 
The differences in the avifaunas between the two 
parcels within the RNI d'Andohahela can be ex- 
plained by two factors: species turnover and geo- 
graphic variation. 

Ninety-three forest-dwelling bird species are 
known from the two parcels. Of these, 78 species 
occur in parcel 1 , of which 46 (59%) are unknown 
from parcel 2. Forty-seven species are document- 
ed in parcel 2, of which 14 species (30%) have 
not been recorded in parcel 1 . This relatively high 
level of species turnover accounts for the biogeo- 
graphic differences between these two sites. These 
93 forest-dwelling biological species correspond 



to 104 phylogenetic species; analyses based on 
phylogenetic species are closely comparable to 
those based on biological species. 

Nonetheless reliance on the biological species 
concept may underestimate levels of genetic dis- 
junctions across the faunal boundary. We strongly 
suspect that genetic studies of taxon pairs (con- 
specific subspecies/phylogenetic species) across 
the ecotone would show that in most cases these 
populations are genetically distinct with little to 
no gene flow between them. For example, Nesillas 
typica formerly was treated as a polytypic biolog- 
ical species, with N. t. typica found in the humid 
areas in southeastern Madagascar and N. t. lantzii 
found in the dry areas to the west. Recent genetic 
studies have demonstrated that these populations 
are distinct and should be treated as separate spe- 
cies (Schulenberg et al., 1993). Several other spe- 
cies might show parallel patterns of differentiation 



Table 17. Faunal similarity indices using a phylogenetic species concept of resident forest-dwelling birds of 
several well-known sites. Above the diagonal is Jaccard's Index and below the diagonal is Simpson's Index. 1 Locality 
abbreviations follow Table 15. 



Anl 



And 



Ran 



Anl 



Anj 



Mas 



Mda 



Anil 



Ber 



Bez 



Zom 



Anl 


— 


0.93 


0.89 


0.84 


0.85 


0.73 


0.51 


0.23 


0.25 


0.22 


0.21 


And 


0.% 


— 


0.86 


0.86 


0.78 


0.71 


0.47 


0.21 


0.33 


0.24 


0.25 


Ran 


0.97 


0.96 


— 


0.87 


0.76 


0.71 


0.56 


0.20 


0.24 


0.24 


0.26 


Anl 


0.92 


0.94 


0.97 


— 


0.80 


0.77 


0.53 


0.20 


0.27 


0.23 


0.25 


Anj 


0.91 


0.88 


0.90 


0.89 


— 


0.81 


0.51 


0.20 


0.21 


0.21 


0.23 


Mas 


0.86 


0.85 


0.88 


0.88 


0.90 


— 


0.58 


0.22 


0.27 


0.25 


0.27 


Mda 


0.83 


0.79 


0.85 


0.87 


0.85 


0.92 


— 


0.27 


0.31 


0.27 


0.29 


Anil 


0.49 


0.47 


0.43 


0.45 


0.45 


0.49 


0.47 


— 


0.84 


0.87 


0.75 


Ber 


0.48 


0.50 


0.45 


0.52 


0.43 


0.52 


0.49 


1.00 


— 


0.86 


0.77 


Bez 


0.46 


0.50 


0.48 


0.48 


0.46 


0.52 


0.44 


0.96 


0.98 


— 


0.78 


Zom 


0.37 


0.47 


0.47 


0.47 


0.46 


0.51 


0.47 


0.94 


0.88 


0.94 


— 



See text (p. 105) for definitions of indices. 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



109 



.02 



.01 



.004 
.02 



.10 
.08 



.04 

— Andohahela (parcel 1) 



Andringitra 
Ranomafana 



.003 
.004 



.007 



07 Analamazaotra 



.01 



.10 



.20 



.04 



.17 



A) 



— — Anjanaharibe-Sud 
Masoala 
Montagne d'Ambre 

.05 

01 I — Andohahela (parcel 2) 
Beza Mahafaly 

06 o 
Berenty 



0.10 



.01 



'• — Ranomafana 

— Analamazaotra 

02 

— : — Andohahela (parcel 1) 

02 

■ Andringitra 



— Anjanaharibe-Sud 



.02 

.02 I Masoala 



.03 



.09 



.01 



.02 



.03 



.11 
.14 



■ Zombitse 

03 

r — Andohahela (parcel 1) 
Andringitra 

— Ranomafana 



B) 



Montagne d'Ambre 

.01 

01 I Andohahela (parcel 2) 

I — Beza Mahafaly 

02 „ 
Berenty 



.02 



.003 

.004 



.03 



^ Analamazaotra 



.02 



.23 



.09 

Anjanaharibe-Sud 

.12 

Masoala 

Montagne d'Ambre 



.19 



.07 

I — Andohahela (parcel 2) 



.04 



.26 



C) 



— Beza Mahafaly 

07 

Berenty 



— — Zombitse 

r — Ranomafana 
— Andohahela (parcel 1) 
Analamazaotra 



02 

- Andringitra 



05 

Anjanaharibe-Sud 



.23 



- Zombitse 



D) 



.04 

03 I Masoala 

I Montagne d'Ambre 

■01 „ 

+— Berenty 

.01 I — Beza Mahafaly 

.02 

1 Andohahela (parcel 2) 



— — Zombitse 



Fig. 17. Cluster analysis of faunal similarity of resident forest birds found at various sites on Madagascar Co- 
ncients derived from Jaccard's and Simpson's (Tables 16, 17) indices were used. Jaccard's Index with biological 
>ecies (A), Simpson's Index with biological species (B), Jaccard's Index with phylogenetic species (C), and Simp- 
m s Index with phylogenetic species (D). See text (p. 105) for definitions of indices. 



110 



FIELDIANA. ZOOLOGY 



(e.g., Agapornis carta, Neomixis striatigula). 
However, this pattern is repeated among the re- 
gional avifauna and among numerous other types 
of vertebrates. 

Despite the current sharpness of fauna! turnover 
at this ecotone, the faunal boundary was not con- 
stant over recent geological time. There have been 
considerable fluctuations in the regional biological 
communities. The only known Holocene subfossil 
site in the region is the Grotte d'Andrahomana, 
located about 50 km west-southwest of Tolagnaro, 
in a region that is now just west of the pluvio- 
metric fault and now largely has a spiny forest 
flora. The site, which was excavated in the early 
portion of this century (Grandidier, 1902; Good- 
man & Rakotondravony, 1996), contained a rich 
assortment of vertebrate material. Among these 
remains were many species of extinct and extant 
lemurs (Walker, 1967), some of which have affin- 
ities to the humid forest or to the dry forest (God- 
frey et al., 1997). The type of Microgale decaryi 
was excavated from this cave (Grandidier, 1928); 
a recent revision of this genus has shown that M. 
decaryi is a synonym of M. principula (MacPhee, 
1987), a species that is widespread in the eastern 
humid forest. Further, remains of an extinct ro- 
dent, Hypogeomys australis, whose sole extant 
congener lives in dry forest, have been excavated 
from the cave (Grandidier, 1903). Radiocarbon 
dating of Hypogeomys remains from the Grotte 
d'Andrahomana show that these animals existed 
in the region 4,440 ± 60 years ago (Goodman & 
Rakotondravony, 1996). 

BlOGEOGRAPHIC AFFINITIES OF THE LOCAL AVI- 
FAUNA in Comparison with Other Well-Known 
Sites on - the Island — On a broader, island-wide 
scale, several other interesting comparisons 
emerged in the biogeographic analysis. Within the 
Jaccard's indices (Figs. 17 A, C) the humid forest 
sites are arranged along a latitudinal gradient from 
south to north. For both biological and phyloge- 
netic species concepts, the distances in the 
branching pattern between the sites of the RNI 
d'Andohahela (parcel 1) north to the RS 
d'Anjanaharibe-Sud are small, which reflects a 
broadly distributed eastern humid forest avifauna. 
On a finer scale, the latitudinal component reflects 
an increase in species richness along a south- 
north gradient. In the Simpson's analyses (Figs. 
17B, D), Montagne d'Ambre and the Masoala 
Peninsula have been placed on a separate branch, 
presumably reflecting a different avifauna! com- 
position. Montagne d'Ambre is an isolated moun- 
tain, with humid forest on the northeastern and 



eastern sides and dry forest along the other flanks. 
The avifauna of the mountain is depauperate in 
comparison with other humid forest sites and pos- 
sesses elements of dry forest. In general, the Ma- 
soala Peninsula has a rich avifauna, and its clus- 
tering with Montagne d'Ambre is presumably re- 
lated to the absence at these two sites of several 
species of broadly distributed eastern humid forest 
birds (Neodrepanis hypoxantha, Dromaeocercus 
brunneus, Cryptosylvicola randrianasoloi, Randia 
pseudozosterops, Hartertula flavoviridis, Crossle- 
yia xanthophrys). 

Within the dry and transitional forest analyses 
for both indices and both species concepts, there 
is a consistent relationship. All of the four sites, 
RNI d'Andohahela (parcel 2), RP de Berenty, RS 
de Beza Mahafaly, and the Zombitse Forest, are 
closely clustered, with short distances between the 
nodes and branches. Although several of these 
sites are separated by considerable distances 
(e.g., RNI d'Andohahela [parcel 2] and the Zom- 
bitse Forest are separated by about 300 km), there 
is a group of species that makes up the regional 
avifauna. Moreover, the birds of Zombitse Forest, 
which is floristically transitional between the east 
and west (Morat, 1973; Du Puy et al., 1994) and 
structurally closer to dry forest, are much more 
closely allied to the dry forest than to the humid 
forest. However, in the dendrograms the Zombitse 
Forest is the outlier of the dry forest sites. 

On the basis of these analyses of the biogeo- 
graphic relationships of the birds known from 
well-documented sites on the island, there appear 
to be at least two distinct avifaunas, one of dry 
areas and one of wet areas. Abiotic factors (e.g., 
annual precipitation) and biotic factors (e.g., flo- 
ristic communities) best explain the separation of 
these two communities. Distance is of little con- 
sequence in explaining this pattern, and sites sep- 
arated by a few kilometers (e.g., parcels 1 and 2 
of the RNI d'Andohahela) have vastly different 
avifaunas. 



Conservation Problems in Southeastern 
Madagascar 

By far the greatest threat to bird populations in 
southeastern Madagascar is posed by habitat de- 
struction, especially of forested habitats. This 
problem affects all forest types throughout the re- 
gion and is driven by a variety of social factors. 
Here we describe in greater detail the history of 
some sites at which we have worked in the region 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



111 






and the forces we have identified that, if uncheck- 
ed, will lead to the loss or degradation of most of 
what little forest remains. 

The Fate of the Remaining Lowland Forest: 
The Case of Analalava and Marovony — Older 
people in the village of Soavary (at the southern 
edge of the Marovony Forest) remember from 
their childhoods, 40-60 years ago, vast stretches 
of forest on the south side of the Ambolomitsaky 
River reaching all the way to Manantenina. In the 
past 50 or so years this forest has been reduced 
to a fraction of its previous size, and now the 
Analalava Forest is a relatively small, isolated for- 
est fragment (Fig. 1 8) — much smaller than shown 
on the map (1:100,000) of the area published in 
June 1961 by the Institut National de Geodesie et 
Cartographic Even during the interval between 
our visits in November 1989 and November 1990 
there was a detectable reduction in the size of the 
Analalava Forest. Most of this change was not a 
result of forest cutting but rather of the encroach- 
ment on the forest of fires in surrounding grass- 
lands (see Salomon, 1993). However, the Maro- 
vony Forest, farther to the north, appears to have 
remained relatively intact. Because the forest 
structure and flora of the Marovony Forest and the 
remaining portion of the Analalava Forest are al- 
most identical (Lowry & Faber-Langendoen, 
1991) and the two forests were until recent times 
essentially continuous, the question can be posed, 
What effect have forest fragmentation and isola- 
tion had on the avifauna of these two areas? 

Approximately equal effort was spent on bird 
surveys at Analalava (13 days) and at Marovony 
(10 days). No migrant species was observed at 
either site, and one species of introduced bird was 
noted at both sites. Nine bird species (eight of 
which are endemic to Madagascar) found in the 
Marovony Forest were not recorded in the Ana- 
lalava Forest, and only one forest-dependent bird 
species (Phyllastrephus zosterops) was found in 
the Analalava Forest and not in the Marovony 
Forest. Given our assumption that the forests once 
had similar avifaunas, and the current differences 
in species richness between these sites, it appears 
that within a short period of time a number of 
local extinctions have occurred in the Analalava 
Forest that are presumably related to its reduction 
in size and increasing isolation. A similar pattern 
has been found in a series of forested fragments 
in the RS d'Ambohitantely (Langrand & Wilme, 
1997). 

Bird densities, as measured by mist-netting suc- 
cess, also were higher in the Marovony Forest 



than in the Analalava Forest (Table 18). When 
only the first 33 net-days of the Marovony Forest 
netting results are used, basically equal to the 
number tabulated from the Analalava Forest, the 
capture rate on average still remains slightly high- 
er than that at the Analalava Forest (31 birds in 
33 net-days = 0.94 capture rate). 

Little remains of the once extensive humid for- 
ests of eastern Madagascar (Green & Sussman, 
1990), particularly the forests below 100 m, and 
very little of such lowland forest is included with- 
in the present reserve system (Nicoll & Langrand, 
1989). Without question the Marovony Forest is 
the largest remaining tract of such forest south of 
Fianarantsoa, and indeed it may be the largest re- 
maining lowland forest south of Maroantsetra. 

During our stay in the Marovony Forest we had 
several discussions with local people about the 
present state of local forests and the effects of 
deforestation. The president of the local commit- 
tee in Soavary observed that the people of this 
community had watched the forests of Analalava 
disappear over the past two generations and that 
they wanted to safeguard the Marovony Forest as 
a managed resource for future generations, spe- 
cifically as a source for important medicinal plants 
and trees for canoes. The president specifically 
asked us about the mechanisms needed and pro- 
cedures to follow to accomplish this. Thus, given 
the biotic importance of the Marovony Forest, 
with regard to the amount of lowland forest within 
the existing reserve system, and the eagerness of 
local people to protect and manage this forest, 
such a project should be considered one of the 
highest priorities for conservation action in south- 
eastern Madagascar. 

The Destruction of the Bezavona (Naham- 
poana) Forest, an Important Watershed for 
tolagnaro, over the course of 3 years — to- 
lagnaro is the largest town in southeastern Mad- 
agascar. The greatest part of the fresh water for 
the city comes from surrounding lakes and moun- 
tain streams. The Bezavona Forest was the water- 
shed for a small stream that in 1989 supplied 12- 
15% of the water used in Tolagnaro (JIRAMA, 
Tolagnaro, pers. comm.). Along one of the middle 
slopes of the Bezavona Forest a concrete em- 
bankment acted as a dam across a small river. A 
pipeline carried water from this reservoir down 
through the forest to the Lakandava water-pro- 
cessing plant and then to Tolagnaro. 

When we first visited the Bezavona Forest in 
November 1989 most of the forest between the 
water station and the dam was in relatively good 






112 



FIELDIANA: ZOOLOGY 




Fig. 18. Remaining portion of the Analalava Forest after vast areas were cleared over the past few decades. 
(Photograph by M. Pidgeon.) 



condition. Between the end of the 1989 and be- 
ginning of the 1990 field seasons, however, a 
number of woodcutters moved into the Bezavona 
Forest. Sections of the forest between the JIRAMA 
station and the dam were partially logged. Trails 
in the nearby forest were expanded, and new trails 



were cut into areas within the Bezavona drainage. 
The acceleration of forest destruction and the 
number of people working the Bezavona Forest 
for wood products was probably related to a re- 
cently imposed ban on woodcutting in the Man- 
dena Forest, a few kilometers away. Traditionally, 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



113 



Table 18. Comparison of bird-netting capture rate in lowland humid forest at Analalava and Marovony. 





Analalava 


Marovony 


Marovony 




(net-day: 


5 = 31) 


(net-days ■ 71) 


(first 33 net-days) 




Total 


Capture 


Total 


Capture 


Total 


Capture 


Species 


captured 


rate 


captured 


rate 


captured 


rate 


Accipiter francesii 






1 


0.01 


1 


0.03 


Caprimulgus enarratus 






1 


0.01 


1 


0.03 


Ispidina madagascariensis 






1 


0.01 


1 


0.03 


Phyllastrephus madagascariensis 


8 


0.26 


18 


0.25 


10 


0.30 


Phyllastrephus zosterops 


1 


0.03 










Hypsipetes madagascariensis 






3 


0.04 


1 


0.03 


Copsychus albospecularis 


2 


0.06 


14 


0.20 


8 


0.24 


Nesillas typica 






2 


0.03 


1 


0.30 


Newtonia amphichroa 






1 


0.01 






Newtonia brunneicauda 


1 


0.03 


1 


0.01 






Terpsiphone mutata 


6 


0.19 


8 


0.11 


5 


0.15 


Oxylabes madagascariensis 






6 


0.08 






Mystacornis crossleyi 






2 


0.03 






Nectarinia souimanga 


1 


0.03 


3 


0.04 


2 


0.06 


Nectarinia notata 


3 


0.10 










Zosterops maderaspatana 


2 


0.06 


3 


0.04 


2 


0.06 


Schetba rufa 


3 


0.10 


1 


0.01 






Dicrurus forficatus 






1 


0.01 






Ploceus nelicourvi 






5 


0.07 






Total numbers 


27 


0.87 


70 


0.99 


31 


0.94 



Mandena provided significant amounts of fuel- 
wood and charcoal for Tolagnaro, and wood re- 
moval and opportunistic hunting have significant- 
ly altered the integrity of this forest (Fig. 19). 
Once restrictions and forest guards were put in 
place at Mandena, woodcutters simply moved 
over to the Bezavona Forest. On several occasions 
officials working for JIRAMA and DEF in Tolag- 
naro were briefed on the destruction of the Be- 
zavona Forest. We were told that forest guards 
had been assigned to the Bezavona Forest. 

When Bezavona was visited on 25 December 
1992, the vast majority of the forest between the 
embankment and the station had been partially to 
totally destroyed. Numerous fires were still smol- 
dering. Furthermore, above the embankment a rel- 
atively flat area of several hectares had been to- 
tally cleared. An attendant at the Lakandava Sta- 
tion mentioned that the quality of the water had 
diminished, that they were having problems with 
sedimentation, and that soon it would be neces- 
sary to relocate the station to another watershed. 
Thus, in the course of 3 years an important forest 
block was lost that represented a large fraction of 
the fresh water for Tolagnaro. 

Two aspects of this situation need to be dis- 
cussed in some detail. The first is that the amount 
of remaining intact forest in the region is small, 
and it will not be a simple matter to find an ap- 



propriate site to relocate the station. Further, such 
a shift is expensive and the money may have been 
better put toward forest protection. The second 
point is that in economic terms it was possible to 
put some clear value to the importance of pro- 
tecting this forest. Without water, Tolagnaro sim- 
ply dries up. The ramifications of this would be 
increased hardships for the local population, the 
economy, and the governmental and private-sec- 
tor infrastructure of southeastern Madagascar. 
Factors more difficult to define, such as biodiver- 
sity, aesthetics, and so forth, do not enter into the 
rationale or justification of why it was important 
to have protected the Bezavona Forest. Alas, of- 
ficials did not extend protection to this forest, and 
it is not clear if, in the long term, any forest in 
the region can be conserved when government of- 
ficials fail to appreciate the value of intact forests 
and do not act to protect them as charged by law. 
The Fate of the Spiny Forest Biome of 
Southern Madagascar — The spiny forest biome 
of southern Madagascar, a forest type with up to 
95% plant endemism (Perrier de la Bathie, 1936), 
is probably the region's most important biome in 
terms of biological uniqueness and endemism. 
Unfortunately, extensive areas of spiny forest 
have been destroyed, from a mixture of commer- 
cial gain for a few and the needs for basic sub- 
sistence for many. 



14 



FIELDIANA: ZOOLOGY 




Fig. 19. Charcoal pit in the Mandena Forest. (Photograph by T. S. Schulenberg.) 



Since the 1920s much of the vegetation of the 
Mandrare River Valley, an area of considerable 
species richness within the subdesert biome, has 
been lost. This process was initiated by several 
colonial families, first for timber and some min- 
ing, and subsequently large tracts were cleared in 
the valley for sisal plantations. The sisal factory 
at Berenty village, near the Malaza Forest, was 
established in 1930 (O'Connor, 1987). The Mal- 
aza Forest was designated as a private reserve in 
1936, when it was fenced and patrolled by guards. 
In the 1980s this reserve became well known as 
the RP de Berenty. Most of the remaining forest 
blocks of gallery vegetation along the lower Man- 
drare River are isolated from each other, are in a 
highly degraded condition, and are surrounded by 
sisal plantations and cleared forest. Although the 
exploitation of spiny forest has been in place for 
decades, new areas are still being converted to 
sisal plantations. Sisal monoculture is insensitive 
to the natural environment (Fig. 15). It is an old 
legacy that shows the devastating and lasting ef- 
fects of indiscriminate exploitation and lack of 
causality associated with the destruction of this 
particularly fragile spiny forest environment. 



Madagascar is a country of approximately 12 
million people, the vast majority of whom depend 
largely on wood and charcoal fuels for household 
use and for small industry. A considerable portion 
of this fuel is obtained from the hardwoods and 
saplings of the spiny forest region of the island. 
Charcoal production and timber exploitation by 
individuals and entrepreneurs is the most serious 
threat to this forest type. 

To put the extent of utilization in perspective, 
95% of household fuel needs in 1990 across Mad- 
agascar were met by wood (FAO, 1993). Up to 
the beginning of the 1980s, 5,000 ha of forest was 
cleared annually to provide Toliara with charcoal 
(Andrianbololona, 1979). Between 1990 and 1991 
fuelwood and charcoal production increased 
446% and 353%, respectively (Office National de 
l'Environnement, 1994). 

Most of the human population of southern 
Madagascar, especially those living in the envi- 
rons of the spiny forest, live in a rural environ- 
ment. Traditionally, exploitation of wood was re- 
stricted to meeting simple daily needs. Much of 
the current destructive practice of clear-felling 
timber to make charcoal was introduced from 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



115 



charcoal workers migrating into the area from the 
regions around Toliara, where they effectively ex- 
hausted the more accessible forests along the ma- 
jor routes. 

Much of the timber exploitation, particularly 
for planks of fantsihilotra {Alluaudia procera) 
and for the production of charcoal, to a varying 
degree is controlled by merchants who contract 
out the labor to local people. The huge increase 
in recent years in charcoal production south of 
parcel 2 of the RNI d' Andohahela is partly a con- 
sequence of demand by entrepreneurs in Toliara 
and other provincial towns for katrafay charcoal. 
Katrafay is a forest hardwood but is also a generic 
term for natural forest charcoal. Such charcoal is 
preferred to kininina (produced from Eucalyptus) 
for its longer and hotter burning properties. 

There are other threats to the spiny forest, and 
most are exacerbated by wood exploitation. Fire 
poses a threat only when colonizing shrubs and 
grasses provide a herbaceous layer in degraded 
forest. In its natural state, the flora of the spiny 
forest, with its succulent characteristics, remains 
largely fire resistant. The several million cattle 
and goats that inhabit the south are destructive to 
the vegetation because of trampling and con- 
sumption of both herbaceous and woody plants on 
a massive scale. 

Today, spiny forest in pristine or relatively in- 
tact condition is perpetually under threat. East of 
the Mandrare River only a few areas within and 
north of parcel 2 of the RNI d' Andohahela show 
the floral and structural heterogeneity of pristine 
spiny forest, and even within these areas it is rare 
to find 60-year-old Alluaudia procera that have 
escaped the pressures of selective logging. Be- 
cause much of the spiny forest is free from large- 
scale burning and intensive agriculture, it is par- 
ticularly saddening to see it pillaged for fuel. Cer- 
tainly the local fuel needs of southeastern Mada- 
gascar and probably further afield could be met 
by the full use of governmental eucalyptus plan- 
tations, which produce a usable charcoal. Anta- 
nanarivo and other major towns on the Central 
High Plateau satisfy their charcoal needs largely 
with eucalyptus. Now is the critical period to put 
in place programs that provide the cultural and 
logistical means to switch charcoal production 
from katrafay to kininina. It is a sad state of af- 
fairs when eucalyptus in the south of the country 
is treated with more reverence and protected by 
stronger laws than is Madagascar's most unique 
floral biome. 



Acknowledgments 

The field studies of SMG and TSS during 1989 
and 1990 in southeastern Madagascar were large- 
ly funded by QIT Madagascar Minerals Ltd. For 
permits to conduct this research we are grateful 
to the Direction des Eaux et Forets, particularly 
Georges Rakotonarivo and Celestine Ravaoari- 
noromanga. Other participants in the QIT-Fer sur- 
veys included G. Ken Creighton, Ron Crombie, 
Patrick Daniels, Claire Hemingway, Ron Nuss- 
baum, Ernestine Raholimavo, Haja Nirina Rako- 
tomanana, Henri Jonah Ratsimbazafy, Chris Rax- 
worthy, Jean Claude Razafimahaimodison, and 
Jim Ryan. Numerous people in the Tolagnaro re- 
gion provided help with permits, logistics, trans- 
portation, etc., but we would particularly like to 
acknowledge Alain Bouffard, Rick Fader, John 
Hatch, Serge LaChapelle, and Andriatsimiova Ra- 
zafindraibe. The 1995 ornithological survey of the 
RNI d' Andohahela was part of a biological in- 
ventory organized by WWF, Madagascar. We are 
grateful to WWF personnel in Tolagnaro and An- 
tananarivo, including Lala Andriamanarivo, Mark 
Fenn, Olivier Langrand, and Sheila O'Connor. 
Joanna Durbin, Brian Fisher, Stig Jensen, Olivier 
Langrand, and Sheila O'Connor provided unpub- 
lished information on the birds of the region. 
Olivier Langrand graciously translated the ab- 
stract into French. 

We are grateful to the curators of the following 
museums for loaning or allowing us access to ma- 
terial under their care: George Barrowclough, Stu- 
art Keith, and Mary LeCroy, American Museum 
of Natural History, New York; Peter Colston, The 
Natural History Museum, formerly the British 
Museum (Natural History), Tring, England; D. 
Stefan Peters, Forschungsinstitut und Naturmu- 
seum Senckenberg, Frankfort; Jean-Francois 
Voisin, Museum National d'Histoire Naturelle, 
Paris; Leon Bennun, National Museum of Kenya, 
Nairobi; Storrs Olson, National Museum of Nat- 
ural History, Washington, D.C.; and Voara Ran- 
drianasolo and Albert Randrianjafy, Pare Bota- 
nique et Zoologique de Tsimbazaza, Antananari- 
vo, Madagascar. 

For the identifications of plants we are grateful 
to George Schatz and Gordon MacPherson, Mis- 
souri Botanical Garden, and Armand Rakotozafy, 
Pare Botanique et Zoologique de Tsimbazaza; for 
insects to Tom Moore, University of Michigan 
Museum of Zoology, and Phil Parrillo, Field Mu- 
seum of Natural History; and for reptiles and am- 
phibians to Chris Raxworthy, University of Mich- 



116 



FIELDIANA: ZOOLOGY 



igan Museum of Zoology. Publication of this 
work was aided by a generous subsidy from QIT 
Madagascar Minerals Ltd. For comments on a 
earlier draft of the manuscript we are grateful to 
Olivier Langrand and Mike Putnam. 



. 1933. In den Urwaldern auf Madagaskar. Die 

Umschau, 37: 30-33. 

[1951]. Katalog der Sammlung von injizierten 



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GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



121 



Appendix 1. Gazetteer of localities mentioned in the text. 





Eastern latitude 


Southern longitude 


Elevation 


Locality 





' 





' 


(m) 


Ambavarano, Lac 


47 


03 


24 


58 




Ambatoabo 


46 


40 


24 


51 




Ambatomaniha, Col d' 


46 


45 


24 


46 


-1000 


Amboasary-Sud 


46 


24 


25 


03 




Ambohitantely, RS 


-47 


16 


-18 


09 


1450-1660 


Ambolomitsaky River 


47 


19 


24 


48 




Ambovombe 


46 


05 


25 


11 




Ampamakiesiny, Col d' 


46 


50 


24 


31 


1375 


Ampanihy 


44 


45 


24 


42 




Amparihy-Est 


47 


21 


23 


57 


30 


Analalava Forest 


47 


19 


24 


13 


20-50 


Analamazaotra, RS 


48 


28 


18 


28 


930-1040 


Andohafasy, River 


47 


12 


24 


47 


0-20 


Andohahela, Pic 


46 


42 


24 


38 


1935 


Andonabe 


46 


49 


24 


41 


-100 


Andrahomana Cave 


46 


40 


25 


50 




Andranohela River 


-46 


47 


-24 


38 




Andraraky Hills 


46 


38 


24 


58 




Andratina River 


46 


07 


24 


34 




Andratoloharano, Lac 


46 


51 


25 


06 


0-10 


Andriambe, Lac 


46 


55 


25 


04 


0-10 


Anjanaharibe-Sud, RS 


-49 


26 


-14 


42 


500-2064 


Anjapolo 


46 


12 


24 


54 


-50 


Ankapaky 


46 


39 


25 


12 




Ankapoky Forest 


46 


31 


24 


59 




Ankarafantsika, RNI 


-46 


57 


-16 


09 


80-330 


Ankepotsy 


46 


43 


24 


32 


1500 


Anony, Lac 


46 


31 


25 


08 


0-10 


Antanandava Forest 


46 


48 


24 


34 




Antseva 


46 


48 


24 


31 


850 


Bealoka 


46 


16 


24 


57 




Beampingaratra 


46 


51 


24 


28 




Befotaka-Sud 


46 


59 


23 


50 


740 


Behara 


46 


23 


24 


57 


40 


Bemangidy 


47 


14 


24 


34 


-100 


Bemangily 


(see Bemangidy) 








Berenty, RP 


46 


17 


24 


59 




Bevala 


46 


24 


25 


06 


-40 


Bevilany 


46 


36 


25 


01 


-100 


Bezavona 


46 


58 


25 


01 




Ebelo 


46 


02 


24 


29 




Efaho River 


46 


52 


25 


48 




Ejeda 


44 


31 


24 


20 




Eminiminy 


46 


49 


24 


41 


-300 


Enakara 


46 


54 


24 


37 




Enaniliha 


46 


53 


24 


39 




Enosiary 


46 


49 


24 


40 




Erombo, Lac 


46 


37 


25 


09 


0-15 


Esira 


46 


43 


24 


20 


400 


Esomony 


46 


38 


24 


30 


530 


Evasia 


46 


42 


24 


46 




Fampanambo 


49 


39 


15 


21 




Fampanombo 


(see Fampanambo) 








Farafangana 


47 


50 


22 


49 




Fenoevo 


46 


53 


24 


42 




Fort-Dauphin 


(see Tolagnaro) 








Fotsivolo 


46 


35 


24 


58 




Hazofotsy 


46 


33 


24 


49 


-100 



122 



FIELDIANA: ZOOLOGY 






Appendix 1. Continued. 



Eastern latitude 



Southern longitude 



Locality 



Elevation 
(m) 



Ifotaka 

Ihotry, Lac 

Imonty 

Isaka-Ivondro 

Isedro, Col de 

Isedro Trail 

Itapera 

Itrafanaomby 

Kirindy Forest 

Lakandava Forest 

Lanirano, Lac 

Lokaro 

Mahamavo 

Mahamavo, Col de 

Malaza Forest 

Manambaro 

Manampanihy River 

Mananafy 

Mananivo, Lac 

Manantantely Forest 

Manantenina 

Mandena 

Mandrare River 

M. in. mar. i River 

Manangotry, Col de 

Mananivo, Lac 

Manombo, RS 

Maroalina 

Maroantsetra 

Marojejy, RNI 

Marosalohy Forest 

Marosohy Forest 

Marosohy, Col de 

Marovony Forest 

Marotsiva 

Mokobe 

Morondava 

Nahampoana 

Namoroka, RNI de 

Petriky 

Pic St. Louis 

Pointe Evatra 

Ranomafana, PN 

Ranomafana Atsimo 

Ranomafana-Sud 

Ranomafana-Tanosy 

Ranomainty 

Ranopiso 

Ranopiso River 

Ranopiso, Col de 

Saihady 

Sedro 

Soavary 

Ste. Luce 

Tanatana, Col de 

Tapera 

Tarantsy River 

Toby 



46 


08 


43 


41 


46 


41 


46 


52 


(see Ambatomaniha, C 


46 


46 


47 


07 


(see Trafonaomby) 


44 


43 


46 


58 


46 


59 


46 


48 


46 


43 


46 


42 


46 


17 


46 


49 


-46 


58 


47 


11 


47 


07 


46 


55 


47 


19 


47 


00 


46 


24 


46 


33 


46 


52 


47 


07 


47 


44 


46 


51 


49 


44 


-49 


15 


46 


51 


46 


49 


46 


48 


47 


20 


46 


47 


46 


38 


44 


17 


46 


58 


-45 


20 


46 


53 


46 


58 


47 


06 


47 


28 


(see Ranomafana-Sud) 


46 


57 


(see Ranomafana-Sud) 


46 


32 


46 


42 


46 


41 


46 


39 


47 


06 


(see Isedro) 




46 


59 


(see Manafiafy) 


46 


51 


(see Itapera) 




46 


34 


(see Bealoka) 





d') 



24 
21 
24 
24 

24 
24 

20 

25 
25 
24 
24 
24 
24 
25 

-27 
24 
24 
24 
24 
24 
25 
24 
24 
24 
23 
24 
15 

-14 
24 
24 
24 
24 
24 
24 
20 
24 

-16 
25 
25 
24 
21 

24 

25 
25 
25 
25 
24 

24 

24 

25 



48 
56 
49 
48 

46 
53 

03 
01 
01 
41 
46 
38 
59 
02 
35 
45 
56 
59 
17 
58 
03 
50 
45 
55 
02 
35 
26 
26 
30 
34 
32 
05 
31 
58 
17 
58 
27 
04 
01 
59 
16 

34 

00 
04 
04 
02 

57 

08 
44 
00 



0-20 



0-20 

0-20 

370 



0-20 

50-600 
30 
0-20 



-830 
0-137 

75-2133 

350-1300 
-1300 
50-100 



75-300 
70-200 

0-40 
530 

0-20 



40 



-300 
-10-40 

-30 

-750 

-70 



GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 






123 



Appendix 1. Continued. 





Eastern latitude 


Southern longitude 


Elevation 


Locality 





' 





' 


(m) 


Tolagnaro 


46 


59 


25 


01 


0-40 


Trafonaomby, Pic 


46 


44 


24 


33 


1956 


Tranomaro 


46 


39 


24 


36 




Tsiombe 


45 


29 


25 


18 




Tsitongambarika, Col de 


47 


00 


24 


42 


-800 


Tsitongatona River 


46 


49 


24 


35 




Tsivory 


46 


05 


24 


04 




Varavara Forest 


46 


43 


24 


30 




Varavara, Col 


46 


43 


24 


31 




Varinadambo 


46 


50 


24 


45 




Vohibaka 


46 


46 


24 


32 




Vohidagaro Hills 


46 


35 


24 


55 


1005 


Vohimainty Hills 


46 


35 


24 


55 


965 


Vohamena, Pic 


46 


58 


24 


43 


1358 


Vohidava Hills 


46 


18 


24 


18 




Vohimena, Pic 


47 


03 


24 


35 


1173 


Vohitsiombe 


46 


09 


24 


26 




Zahamena, RNI 


-48 


50 


-17 


40 


750-1512 



124 



FIELDIANA: ZOOLOGY 



Appendix 2. Names of plant genera, species, and families mentioned in the 
text. 1 



Genus/species 



Family 



Genus/species 



Family 



Abrus 

Acacia farnessiana 

Acacia minnutiflora 

Acacia rovumae 

Acacia sakalava 

Adansonia za 

Agave rigida 

Aguaria 

Alberta 

Albizia polyphylla 

Alluaudia ascendens 

A I Inn ml iii comosa 

Alluaudia humbertii 

Alluaudia procera 

Aloe divaricata 

Aloe vaombe 

Aloe vaotsanda 

Antirohea 

Aphloia theiformes 

Asplenium 

Azima tetracantha 

Bauhinia 

Bulbophyllum 

Canarium obovatum 

Capparis chrysomeia 

Capparis seppiaria 

Cassia 

Casuarina 

Celtis gomphophyla 

Celtis phillipensis 

Cerbera venenifera 

Chrysophyllum boivinianum 

Citrus 

Commelina ramulosa 

Commiphora 

Cordia ronnii 

Cordia sinensis 

Crateva excelsa 

Crotalaria 

Croton 

Croton monge 

Cyathea 

Cynanchum 

Delonix regia 

Dicotna 

Dicoryphe viticoides 

Didierea 

Dilobeia thouarsii 

Diospyros myriophylla 

Dombeya 

Dracaena reflexa 

Dypsis decaryi 

Elaeocarpus 

Erythrina 

Erythroxylum gerrardi 

Eucalyptus citriodora 

Euphorbia leucodendrum 

Euphorbia stenoclada 

Euphorbia tirucalli 

Fernandoa madagascariensis 



Fabaceae 

Fabaceae 

Fabaceae 

Fabaceae 

Fabaceae 

Bombacaceae 

Agavaceae 

Ericaceae 

Rubiaceae 

Fabaceae 

Didiereaceae 

Didiereaceae 

Didiereaceae 

Didiereaceae 

Liliaceae 

Liliaceae 

Liliaceae 

Rubiaceae 

Flacourtiaceae 

Aspleniaceae 

Salvadoraceae 

Leguminosae 

Orchidaceae 

Burseraceae 

Capparaceae 

Capparaceae 

Fabaceae 

Casuarinaceae 

Ulmaceae 

Ulmaceae 

Apocynaceae 

Sapotaceae 

Rutaceae 

Commelinaceae 

Burseraceae 

Boraginaceae 

Boraginaceae 

Capparidaceae 

Leguminosae 

Euphorbiaceae 

Euphorbiaceae 

Cyathaceae 

Asclepiadaceae 

Leguminosae 

Asteraceae 

Hamamelidaceae 

Didiereaceae 

Proteaceae 

Edenaceae 

Sterculiaceae 

Dracaenaceae 

Arecaceae 

Elaeocarpaceae 

Leguminosae 

Erythroxylaceae 

Myrtaceae 

Euphorbiaceae 

Euphorbiaceae 

Euphorbiaceae 

Bignoniaceae 



Ficus grevei 

Ficus megapoda 

Flacou rtia lucidiaefolia 

Gaertnera 

Gyrocarpus americanus 

Hibiscus 

Hippocratea rubignosa 

Humbertia madagascariensis 

Ilex mitis 

Impatiens 

Kalanchoe beharensis 

Kalanchoe gastonis 

Macaranga 

Maeura filiformis 

Malva 

Ma run in fraxineae 

Mascarenhasia 

Medinilla 

Melia azedarach 

Millettia 

Moringa 

Morus 

Nastus 

Neotina isoneura 

Nepenthes madagascariensis 

Nymphaea 

Ocotea 

Oncostemon 

Opuntia 

Pachypodium 

Pandanus spp. 

Philippia 

Phragmites communis 

Phyllanthus seyrigii 

Pithecellobium dulce 

Pittosporum 

Pothos scandens 

Quivisianthe papinae 

Ravenala madagascariensis 

Ravensara 

Rinoria greveana 

Sarcolaena multiflora 

Sarcostemma decorsei 

Sloanea rhodantha 

Sorindeia madagascariensis 

Strongylodon 

Symphonia 

Syzygium 

Tabernaemontana 

Tamarindus indica 

Tambourissa 

Tarenna purinosum 

Tina isoneura 

Typhonodorum 

Uapaca 

Vaccinium 

Vepris sclerophylla 

Weinmannia 

Xerophyta 



Moraceae 

Moraceae 

Flacourtiaceae 

Rubiaceae 

Hernandiaceae 

Malvaceae 

Celastraceae 

Convolvulaccae 

Aquifoliaceae 

Balsaminaceae 

Crassulaceae 

Crassulaceae 

Euphorbiaceae 

Capparidaceae 

Malvaceae 

Marattiaceae 

Apocynaceae 

Mclastomataceae 

Meliaceae 

Leguminosae 

Moringaceae 

Moraceae 

Poaceae 

Sapindaceae 

Nepenthaceae 

Nymphaeaceae 

Lauraceae 

Myrsinaceae 

Cactaceae 

Apocynaceae 

Pandanaceae 

Ericaceae 

Poaceae 

Euphorbiaceae 

Leguminosae 

Pittosporaceae 

Araceae 

Meliaceae 

Strelitziaceae 

Lauraceae 

Vitaceae 

Sarcolaenaceae 

Asclepiadaceae 

Elaeocarpaceae 

Anacardiaceae 

Fabaceae 

Clusiaceae 

Myrtaceae 

Apocynacae 

Fabaceae 

Monimiaceae 

Rubiaceae 

Sapindaceae 

Araceae 

Euphorbiaceae 

Ericaceae 

Rutaceae 

Cunoniaceae 

Velloziaceae 



1 Family allocation generally after Mabberley (1989). 
GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 



125 



Index to Malagasy Vernacular Bird Names 



akanga 34 
akoala 27 
akoholahin 'ala 27 
akohon'ala 27 
aliotsy 47 
andreabokia 71 
angetry 75 
angoiky 79 
aoAa 46 

arakarandroka 39 
aj/iy 60 
bemaso 79 
bevorotse 32 
o/fcAo 84 
bintitra 56 
o/r/Av 36 
o/fa 79 
fc/rra*}' 78 
fcofza 43 
bohaky 42 
bokazava 64 
bokazavo 64 
ooAy 29 
boloky 44 
borisy 62 
or/a 62 
oV/«o 42 
eoAa 46 
fandikalala 49 
fandraokibo 32 
fantsasatry 58 
farivaza 45 
fatsatsatry 11 
fifiokala 82 
Ji/i/aAa 30 
fileliakondro 30 
fililiotra 54, 63 
fitadroanga 68 
fitatr'ala 67 
yj/a/ra 67 
yi/arry 24, 67 
fodibeotse 89 
fodimanta 90 
fodisiay 88 
/<w/y a/a 88 
/oa>' m^na 89 
/o// manga 43 
/o/y manga 43 
folymena 89 
fonomavo 42 
fonomity 43 
goaka 87 
goaAy 87 
goapakala 54 
ZiaAa 51 
hankana 53 
hatrakatra 41 
natrakatraka 41 



/idto /i^Ao 51 

hindry 32 

hitsikitsika 33 

hitsikitsiky 33 

hondria 32 

/jora 51 

horova 66 

horovana 66 

jorioke 62 

kanaka 45 

kariaky 45 

Aar/a 79 

A<»/»o 25 

A/a 43, 44 

A/oo 35 

A/Ay 35 

kimokimo 42 

kirikirioky 57 

A/r/o 57 

kitreoky 45 

AoaAy 87 

kolokoloky 40 

kopaky 54 

kotohake 51 

langopaka 54 

langopaka tataro 54 

lombokoma 25 

/ova 86 

mandom bokomana 25 

maritay 88 

mavoloha 64 

ongongo 28 

papango 30 

parataka 69 

parataky 65 

peeda 67 

paAa/a 47, 58 

pretaka 65 

pretaky 65 

ragado 26 

railovy 86 

ramaro 88 

rangado 26 

razamboay 24 

rehitriky 33 

rengetry 75 

revitsy 56 

revitsy'ala 56 

rimaly 75 

rimaro 88 

sadakely 29 

sa/a/y AWy 38 

.sa/a/y 6^ 38 

sama 28 

sarivazy 45 

simitsy 69 

5/ay 79 

5oy 61, 78 



soy lehibe 79 
soy manga 78 
soy AWy 78 

taitoaky 41 

takatry 26 

takatsy 26 

taleva 36 

talevana 37 

taotaokafo 45 

tatarao-ala 54 

tatarobobaky na lakopaky 58 

teraratsy 57 

^50 50 

retro 50 

te fso manga 50 

r/YoAa 50 

toloho 51 

foro 52 

torotoroka 52 

traotrao 34 

treo-treo 59 

triotrio 63 

tsikaloto 42 

tsikinitry 90 

tsikiriokirioke 57 

tsikodara 59 

tsikoja 36 

tsikonina 66 

tsikorova 66 

tsiksoysoy 78 

tsilaka 67 

tsilotsilon'aka 54, 63 

tsilovanga 82 

tsimala 30 

tsimimitraka 73 

tsimimitrala 74 

tsimimitry 1 1 

tsimimitsy 71 

tsimitsy 73 

tsimpiritry 90 

tsipara 32 

tsipara korovana 32 

tsipiritsy 90 

tsiraraka 57 

tsirikititio 64 

tsiriry 28 

van#a 82 

vaza 44 

vazambe 43 

vazatsihotsy 44 

v/nwy 56 

vorokotsy 25 

vorompijfkoka 84 

vorompotsy 25 

vorondolo 51, 52 

vorondreo 59 

voronzaza 83 

zea-zea 73 



INDEX TO MALAGASY BIRD NAMES 



127 



Index to Scientific Names 



Page numbers in bold indicate the main species account. 



Abrus 42, 125 

Acacia 13, 14, 31, 43, 44, 46, 79, 
86 

farnessiana 44, 45, 77, 125 

minnutiflora 125 

rovumae 14, 24, 30, 31, 32, 33, 45, 48, 57, 73, 77, 
79, 82, 125 

sakalava 11, 125 
Accipiter 30 

francesii 86, 92, 96, 99, 100, 101, 104, 1 14 

francesii francesii 31-32, 106 

henstii 31, 92, 96, 106 

madagascariensis 31, 32, 67, 92, 104, 106 
Accipitridae 29 
Acrantophis dumerilii 46 

Acridotheres tristis 31, 87-88, 94, 97, 104, 105 
Acrocephalus newtoni 68, 93, 96 
Actitis hypoleucos 39, 92, 96 
Actophilornis albinucha 37 
Adansonia 2, 13, 30, 83 

za 5, 88, 125 
Agapornis cana 57, 93, 95, 99, 104, 111 

cana ablactanea 44-45, 106 

cana cana 44—45, 106 
Agave rigida 44, 45, 77, 79, 102, 125 
Aguaria 11, 1 25 
Alaudidae 62 
Alberta 11, 125 
Albizia 13, 86 

polyphylla 14, 30, 45, 48, 49, 77, 125 
Alcedinidae 55 
Alcedo vintsioides 93, 96, 99, 101, 102 

vintsioides vintsioides 55-56 
Alectroenas 43, 93 

madagascariensis 21, 42-43, 96, 106 
Alluaudia 13, 14, 32, 33, 44, 83, 84, 116 

ascendens 5, 15, 125 

comosa 5, 125 

humbertii 13, 125 

procera 5, 44, 45, 82, 84, 116, 125 
Aloe 13, 77 

divaricata 11, 78, 79, 125 

vaombell, 78, 79, 125 

vaotsanda 5, 125 
Anas bernieri 21 

erythrorhyncha 28-29, 92, 96 

hottentota 29, 92 

melleril%, 91, 92 
Anatidae 28 
Antirohea 9, 125 
Anseriformes 28 
Aphloia theiformes 9, 125 
Apodidae 54 
Apodiformes 54 
Apus barbatus [balstoni] 54, 55, 63, 93, 97 

melba [willsi] 55, 63, 93, 97 
Ardea cinerea 92, 96 

cinerea firasa 26 

humbloti 26, 92 

purpurea 92, 96 



purpurea madgascariensis 25-26 
Ardeidae 24 
Ardeola idae 24, 92, 96 

ralloides 24, 92, 96 
Arenaria interpres 39, 92 
Asio capensis [nova] 53, 93 

madagascariensis 52-53, 91, 93, 95, 96, 106 
Asplenium 9, 125 
Atelornis crossleyi 58, 93, 96, 107 

pittioides 58, 91, 93, 96, 102, 107 
Avahi 53 

Aviceda madagascariensis 29, 92, 96, 106 
Azima tetracantha 75, 125 

Bakerella 61, 78, 90 

Bauhinia 14, 125 

Boophis madagascariensis 53 

Brachypteracias leptosomus 57-58, 91, 93, 97, 101, 106 

squamiger 57, 58, 91, 93, 97, 102, 107 
Brachypteraciidae 57, 105 
Bubulcus ibis 24-25, 30, 92, 96 
Bulbophyllum 9, 125 
Bulweria fallax 23 
Buteo 87 

brachypterus 30, 32, 92, 96, 104, 106 
Butorides striatus 92, 96 

striatus rutenbergi 25 

Calicalicus madagascariensis 31, 80, 94, 97, 101, 107 
Calidris alba 39, 92 

ferruginea 39-40, 93 
Calumma nasutus 50, 52, 56 
Campephagidae 63 
Canarium 1 1 

obovatum 9, 125 
Canirallus kioloides 35, 92, 96 

kioloides kioloides 36, 106 
Capparis 66, 72 

chrysomeia 11, 79, 125 

seppiaria 75, 125 
Caprimulgidae 53 
Caprimulgiformes 53 
Caprimulgus enarratus 54, 91, 93, 95, 96, 101, 106, 1 14 

madagascariensis 93, 96, 99, 101 

madagascariensis madagascariensis 53-54 
Cassia 125 
Casuarina 8, 87, 125 
Catharacta antarctica 40 
Celtis gomphophyla 14, 43, 125 

phillipensis 14, 43, 44, 49, 66, 82, 125 
Centropus grillii 50 

toulou 53, 93, 96, 100, 104, 105 

toulou toulou 50-51, 106 
Charadriidae 37 
Charadriiformes 37 
Charadrius hiaticula [tundrae] 38, 92 

leschenaultii 38, 92 

marginatus tenellus 38 

pecuarius 92 

pecuarius pecuarius 38 



128 



FIELDIANA: ZOOLOGY 






thoracicus [bifrontatus] 38, 92, 95 

tricollaris 38, 92 
Chlidonias hybridus [sclateri] 40 
Chrysophyllum boivinianum 9, 11, 1 25 
Ciconiidae 26 
Ciconiiformes 24 
Circus maillardi 30 

Cisticola cherina 70, 94, 97, 99, 104, 105 
Or™ 77, 125 
Collocalia francica 54 
Columbidae 41 
Columbiformes 41 
Commelina ramulosa 12, 13, 125 
Commiphora 13, 125 

Copsxchus albospecularis 31, 93, 97, 99. 100, 101, 102, 
i04, 114 

albospecularis albospecularis 107 

albospecularis inexpectatus 67, 107 

albospecularis pica 67, 1 07 
Coraciidae 57 
Coraciiformes 55 
Coracina cinerea 79, 93, 97, 100 

cinerea cinerea 63-64, 107 

cinerea pallida 63-64, 1 07 
Coracopsis 57 

nigra 93, 96, 100, 104 

nigra libs 43, 106 

nigra nigra 43—44, 106 

vasa 93, 96, 104 

vasa drouhardi 43, 106 

vasa vasa 43, 106 
Cordia 11 

ronnii 66, 79, 125 

sinensis 125 
Corvus albus 82, 86-87, 88, 
Coturnix coturnix [africana] 
Coua 95 

caerulea 48, 49-50, 93, 96, 100, 106 

cristate 93, 96, 97, 104 

cristata cristata 106 

cristata maxima 47-49 

cristata pyropvga 47-49, 1 06 

cursor 47, 93,' 96, 106 

g/gas 45-46, 75, 90, 91, 93, 95, 96, 100, 104, 106 

reynaudii 46-47, 48, 93, 96, 106 

ruficeps 93, 96 

ruficeps olivaceiceps 47, 1 06 

serriana 101, 106 

verreauxi 49 
Crateva ^cc/sa 14, 45, 66, 79, 82, 83, 125 
Crossleyia xanthophrxs 76-77, 94, 97, 101, 102, 107, 

111 
Crotalaria 13, 125 
Croton 12, 13,44,45, 125 

monge 11, 1 25 
Cryptosvlvicola randrianasoloi 68-69, 81, 94, 95, 97, 

107, 111 
Cuculidae 45 
Cuculiformes 45 
Cuculus audeberti 106 

cristata 48 

rochii 45, 93, 96, 106 
Cyanolanius madagascarinus 31, 79, 94, 97, 102 

madagascarinus madagascarinus 84—85, 108 
Cynanchum 13, 77, 125 



94, 97, 104 
34,92 



Cypsiurus par\>us 54-55, 63, 93, 97 
Cythea 11, 125 

Daption capense 23 
Decaryia 13 
Delonix 13 

regia 11, 125 
Dendrocygna bicolor 28, 92, 96 

viduata 28. 92, 96 
Dicoma 11, 125 
Dicoryphe viticoides 11, 1 25 
Dicruridae 86 

Dicrurus forftcatus 30, 79, 81, 87, 94, 97, 99. 100, 101, 
102, 104, 105, 114 

forftcatus forftcatus 86, 108 
Didierea 125 
Dilobeia thouarsii 9, 1 25 
Diomedea cauta 22 

chlororhynchos 23 
Diomedeidae 22 
Diospyros 65 

myriophylla 13, 125 
Dombeya9, 11, 125 
Dracaena reflexa 9, 125 
Dromaeocercus brunneus 70, 94, 97, 107, 111 

seebohmi 107 
Dryolimnas cuvieri 92, 96, 100 

cuvieri cuvieri 35-36, 106 
Dypsis decaryi 12, 13, 55, 125 

Egretta alba 92, 96 

alba melanorhynchos 25 

ardesiaca 25, 92, 96 

dimorpha 25, 92, 96 
Elaeocarpus 11, 1 25 
Eliurus minor 53 

webbi 53 
Erythrina 125 

Erythroxylum gerrardi 12, 125 
Estrildidae 90 
Eucalyptus 16, 31, 116, 125 

citriodora 11 
Eulemur fulvus 43, 50 
Euphorbia 13, 14, 70, 71, 82 

leucodendrum 125 

stenoclada 5, 125 

tirucalli 64, 11, 125 
Euryceros prevostii 101, 1 08 
Eurylamidae 59 
Eurystomus 30, 53, 57, 86 

glaucurus 44, 93, 97, 104 

glaucurus glaucurus 57, 106 
Eutriorchis astur 101, 1 06 

Fa/co concolor 33, 57, 92, 96, 104, 105 

eleonorae 33, 92, 96, 104 

newtoni 30, 92, 96. 99, 104 

newtoni newtoni 32—33, 106 

peregrinus [radama] 33-34, 92, 104, 106 

zoniventris 33. 92. 95, 96, 104, 106 
Falconidae 32 
Falconiformes 29 

Falculea palliata 83, 84, 94, 97, 108 
Fernandoa madagascariensis 64, 77, 78, 79, 125 



INDEX TO SCIENTIFIC NAMES 



129 



Ficus 9, 14, 42, 43, 44 

grevei 42, 82, 125 

megapoda 43, 125 
Flacourtia lucidiaefolia 13, 125 

Fow<//a madagascariensis 33, 87, 89, 94, 97, 99, 101, 
104, 105, 108 

omissa 89-90, 94, 97, 101, 102, 108 
Fulica cristata 37 

Gaertnera 11, 61, 125 

Galidia elegans 82 

Galliformes 34 

Gallinago macrodactyla 39, 92 

Gallinula chloropus 36, 92, 96 

Geogale aurita 51 

Glareola ocularis 37, 92 

Glareolidae 37 

Gruiformes 34 

Gyrocarpus americanus 88, 125 

Hapalemur 53 

Hartertula flavoviridis 74, 94, 97, 101, 102, 107, 111 

Hartlaubius auratus 87, 94, 97, 108 

Heterixalus boettgeri 50 

Hibiscus 72, 77, 125 

Hippocratea rubignosa 44, 125 

Hipposideros commersoni 53 

Hirundinidae 62 

Hirundo rustica 63, 93 

Humbertia madagascariensis 44, 125 

Hypositta corallirostris 85, 94, 97, 107 

perdita 85 
Hypsipetes madagascariensis 3 1 , 53, 79, 86, 93, 97, 99, 
100, 101, 102, 104, 105, 114 

madagascariensis madagascariensis 66-67, 107 
Hypogeomys australis 1 1 1 

Ilex mitis 9, 125 

Impatiens 11, 1 25 

Ispidina madagascariensis 93, 97, 99, 101, 102, 114 

madagascariensis diluta 106 

madagascariensis madagascariensis 56, 106 
Ixobrychus minutus 24, 92 

Jacanidae 37 

Kalanchoe 13 
beharensis 11, 78, 125 
gastonis 125 

Laridae 40 

Larus dominicus 40, 93 

Lemur catta 30, 32, 78 

Leptopterus chabert 94, 97, 99, 104 

chabert chabert 84, 108 

chabert schistocercus 84, 108 

virWw 79, 81, 83, 94, 97, 102, 104 

viridis annae 108 

viridis viridis 83-84, 108 
Leptosomatidae 58 
Leptosomus discolor 58-59, 86, 93, 97 

discolor discolor 107 
Limosa lapponica 92 

lapponica lapponica 38 
Lonchura nana 87, 90, 94, 97, 104 
Lophotibis cristata 91, 92, 95, 96, 100 



cristata cristata 26-27, 106 
cristata urschi 27, 106 
Lygodactylus 31 

Maba 12 

Mabuya 31, 49 

Macaranga 9, 11, 44, 79, 125 

Machaeramphus alcinus [anderssoni] 29, 92, 106 

Maeura filiformis 11, 1 25 

Malva 11, 125 

Mantidactylus lugubris 56 

Marattia fraxineae 11, 1 25 

Margaroperdix madagascarensis 34, 92, 96 

Mascarenhasia 11, 125 

Medinilla 61, 125 

Melia azedarach 44, 66, 125 

Meropidae 56 

Merops superciliosus 93, 91, 100, 101, 104 

superciliosus superciliosus 56-57 
Mesitornis unicolor 34-35, 91, 92, 106 
Mesitornithidae 34 
Microcebus 53 

murinus 51 

rw/ws 81 
Microgale decaryi 1 1 1 

principula 1 1 1 
Millettia 12, 125 
Milvus migrans 92, 96, 104 

migrans parasitus 29-30 
A//ra/ra /low* 62, 93, 97, 104 
Monarchidae 74 
Moringa 13 125 
Moras 42, 125 

Motacilla flaviventris 63, 93, 97, 102 
Motacillidae 63 
Mus musculus 53 
Mycteria ibis 26 
Mystacornis crossleyi 77, 94, 97, 101, 102, 107, 1 14 

Nastus9, 125 
Nectarinia 61, 105 

ziotata 67, 77, 80, 94, 97, 99, 100, 101, 114 

notata notata 78-79, 107 

souimanga 61, 67, 71, 80, 81, 94, 97, 99, 100, 101, 
102, 104, 105, 114 

souimanga apolis 77-78, 107 

souimanga souimanga 77-78, 107 
Nectariniidae 77 
Neodrepanis coruscans 60-61, 91, 93, 97, 101, 102, 107 

hypoxantha 61-62, 78, 93, 97, 102, 107, 111 
Neomixis 81 

striatigula 72, 94, 97, 99, 104, 111 

striatigula pallidior 73, 1 07 

striatigula scalteri 73, 107 

striatigula striatigula 73 

tenella 45, 71, 72-73, 94, 97, 99, 100, 104, 105 

tenella debilis 72, 107 

tenella orientalis 72, 107 

tenella tenella 72, 107 

viridis 72, 73, 94, 97, 107 

viridis viridis 73 
Neotina 86 

isoneura 14, 30, 32, 43, 44, 45, 57, 66, 84, 125 
Nepenthes madagascariensis 14, 125 
Nephila 66, 74, 81 
Nesillas lanztii 69, 94, 97, 107, 109 



130 



FIELDIANA: ZOOLOGY 



typica 94, 97, 101, 102, 109, 114 

typica typica 69, 107, 109 
Newtonia 71, 81 

amphichroa 70-71, 72, 94, 97, 101, 102, 107, 114 

archboldi 71-72, 94, 95, 97, 99, 107 

brunneicauda 32, 70, 72, 79, 81, 94, 97, 99, 100, 101, 
102, 104, 114 

brunneicauda brunneicauda 71, 107 

fanovanae 72, 91, 94, 97, 101, 107 
Nesomys audeberti 76 
Ninox superciliaris 52, 91, 93, 96, 106 
Numenius arquata 39 

phaeopus 39, 92 
Numida meleagris 92, 96 

meleagris mitrata 34 
Numididae 34 

Nycticorax nycticorax 24, 25, 92, 96 
Nymphaea 14, 125 

Ocotea 9, 125 

Oena capensis 41, 93, 96, 97, 99, 104 

capensis aliena A2, 106 
Oncostemon 1 1 , 60, 1 25 
Opiums 33 
Opuntia 11, 79, 125 
Oriolia bernieri 82, 83, 101, 108 
Otus rutilus 53, 93, 96, 99, 101, 102 

rutilus rutilus 51-52, 106 
Oxvlabes madagascariensis 75-76, 81, 94, 97, 101, 102, 
107, 114 

Pachypodium 13, 125 
Pachyptila vittata 23 
Pandanus 8, 14, 27, 125 
Pandion haliaetus 29 
Pandionidae 29 
Passeriformes 59 
Pelecaniformes 23 
Phaethon aethereus 23-24 
Phaethonidae 23 
Phalacrocoracidae 24 
Phalacrocorax africanus 92, 96 

africanus pictilis 23 
Phasianidae 34 
Phedina borbonica 55, 63, 93, 97 

borbonica madagascariensis 62-63 
Phelsuma 31, 81 
Philepitta 61 

castanea 59-60, 91, 93, 97, 101, 102, 107 

schlegeli 21 
Philippia 11, 82, 125 
Phoenicopteridae 27 
Phoenicopterus minor 28 

ruber [roseus] 27-28, 92 
Phragmites 36, 90 

communis 50, 125 
Phyllanthus 9 

seyrigii 42, 79, 1 25 
Phyllastrephus apperti 107 

cinereiceps 64, 65, 66, 93, 97, 102, 107 

madagascariensis 74, 81, 93, 97, 101, 102, 1 14 

madagascariensis inceleber 107 

madagascariensis madagascariensis 64-65, 107 

tenebrosus 101, 107 

zosterops 64, 74, 76, 93, 97, 101, 102, 112, 114 

zosterops andapae 107 



zosterops fulvescens 107 

zosterops maroantsetrae 107 

zosterops zosterops 65-66, 107 
Pithecellobium dulce 65, 75, 81, 82, 87, 125 
Pittosporum 11, 1 25 
Platypelis grandis 53 
Ploceidae 88 
Ploceus nelicourvi 88, 97, 101, 102, 108, 114 

sakalava 30, 33, 67, 94, 97, 99, 104 

sakalava minor 88-89, 94, 108 
Pluvialis squatarola 37 
Podicipedidae 23 
Podicipediformes 23 
Polyboroides 87 

radiatus 30, 92, 96, 106 
Porphyrio porphyrio [madagascariensis] 37, 92 
Porphyrula alleni 37 
Pothos scandens 9, 125 
Procellaridae 23 
Procellariiformes 22 
Propithecus verreauxi 30, 32 
Pseudobias wardi 21, 74, 94, -97, 107 
Pseudocossyphus bensoni 107 

imerinus 68 

j/uir/x>j 93, 97, 102 

sharpei sharpei 68, 107 
Psittacidae 43 
Psittaciformes 43 

Pterocles personatus 41, 93, 96, 106 
Pteroclididae 41 
Pterodroma baraui 23 

macroptera 23 

mollis 23 
Pteropus rufus 30 
Puffinus atrodorsalis 23 

pacificus 23 
Pycnonotidae 64 

Quivisianthe 44 
papinae 30, 43, 45, 125 

Rallidae 35 

Rallus madagascariensis 35 

AW/a pseudozosterops 70, 91, 94, 95, 97, 107, 111 

/?am« 53 

raffus 51 
Ravenala 27, 60 

madagascariensis 9, 14, 17,44, 125 
Ravensara 11, 1 25 
Rinoria greveana 14, 66, 75, 79, 125 
Riparia paludicola 62, 93, 97 
Rostratula benghalensis [benghalensis] 37, 92 
Rostratulidae 37 

Sarcolaena multiflora 44, 125 
Sarcostemma 13, 71 

decorsei 125 
Sarkidiornis melanotos 28, 92, 96 
Sarothrura insularis 36, 91, 92, 96, 106 
Saxicola torquata 93, 97 

torquata sibilla 67-68 
Sc/m?/&j r«/a 85, 94, 97, 101, 1 14 

rw/o occidentalis 108 

rw/a m/a 80-81, 107 
Scolopacidae 38 
Scopidae 26 



INDEX TO SCIENTIFIC NAMES 



131 



Scopus umbretta 92, 96 

umbretta umbretta 26 
Sloanea 10, 41, 61, 89, 90 

rhodantha% 10, 11,49, 125 
Sorindeia madagascariensis 9, 125 
Sphecius grandidieri 33 
Stercorariidae 40 
Stercorarius longicaudus 40 
Sterna bengalensis 41, 93 

bergii 40-41, 93 

caspia 40, 93 

dougallii 40, 93 

hirundo 40 

sandvicensis 40 
Sternidae 40 

Streptopelia picturata 32, 34, 93, 96, 99, 100, 101, 102, 
104 

picturata picturata 41-42, 106 
Strigidae 51 
Strigiformes 51 
Strongylodon 11, 90, 125 
Sturnidae 87 
Sylviidae 68 

Symphonia AA, 50, 90, 125 
Syzygium 9, 125 

Tabernaemontana 14, 125 

Tachybaptus pelzelnii 23 

Tamarindus 14, 30, 32, 43, 44, 46, 59, 64, 77, 86 

indica 14, 45, 57, 75, 77, 79, 125 
Tambourissa 9, 11, 1 25 
Tarenna purinosum 13, 125 
Terpsiphone 75 

mutata 31, 45, 79, 94, 97, 99, 100, 101, 102, 104, 1 14 

mutata mutata 74-75, 107 
Thamnornis chloropetoides 70, 94, 97, 107 
Threskiornithidae 26 
Timaliidae 75 
Tina isoneura 11, 125 
Tracheloptychus 31 



Treron australis 43, 93, 96, 99, 100 

australis australis 42, 106 
Tringa nebularia 39, 92, 96 
Turdidae 67 
Turnicidae 35 

Turnix nigricollis 35, 92, 96, 106 
Tylas eduardi 31, 82, 94, 97, 102 

eduardi eduardi 85-86, 108 
Typhonodorum 14, 125 
Tyto alba 93 

alba affinis 51 

soumagnei 106 
Tytonidae 51 

Uapaca 42, 43, 44, 125 
Upupa epops 93, 97, 99, 104 
epops marginatus 59, 107 
Upupidae 59 
Uroplatus sikorae 53 
Usnea 9, 125 

Vaccinium 11, 125 

Vanga curvirostris 30, 86, 87, 94, 97, 99, 100, 104 

curvirostris cetera 81-82, 108 

curvirostris curvirostris 81-82, 108 
Vangidae 80 
Vepris sclerophylla 13, 125 

Weinmannia 9, 125 

Xenopirostris polleni 82-83, 94, 97, 101, 108 

xenopirostris 82, 94, 97, 108 
Xerophyta 13,41, 125 

Zoonavena grandidieri 63, 96 

grandidieri [grandidieri] 54, 93, 106 
Zosteropidae 79 

Zosterops maderaspatana 31, 80, 94, 97, 101, 102, 104, 
105, 114 

maderaspatana maderaspatana 79, 107 



132 



QA5 82/99 III 

^38956 - ** 



FIELDIANA: ZOOLOGY 




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