UNIVLR^ OF ILLINOIS RY flUBBANA 1PJUQN BIC ,ol yvu n ^or rxo.'SS' i^sieA* bock d»\)er oolosrv The Birds of Southeastern Mada: ar A No von Public hulenberg PUBi I c perinit' e or fra< gments" (if any) and "Literature (periods are not used after abbrci recent issues ot Fietdiana. Chicago Manual of Stvic ( 13th ed.), pu \bbreviations, R< in the following fot Vol. 2, Ti lion. Wash Journal of Ecol n visions, pp. 6 ulJ. whei Illiisirat F1ELDIANA Zoology NEW SERIES, NO. 87 The Birds of Southeastern Madagascar Steven M. Goodman1 2 Mark Pidgeon2 A. F. A. Hawkins3 Thomas S. Schulenberg1 4 department of Zoology Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 USA 2World 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 km2, 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 km2, 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 km2 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 V6 [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 km2 of the 7,000 km2 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 0 0 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 0 July August September October 2 0 5 0 8 11 14 7 November 3 8 December 0 7 1985 January February March 0 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 (<$:$) visited1 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 0 Marosohy 425 m 50 7 3 0 725 m 63 4 12 0 Marovony 50 m 110 16 0 0 Manantantely 100 m 32 7 0 0 Andohahela 440 m 50 8 4 0 810 m 50 1 3 0 1200 m 50 3 7 1 1500 m 50 0 0 4 1875 m 50 0 0 0 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 locations3 Sta- tus1 Habi- tat2 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 locations3 Sta- tus' Habi- tat2 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 locations3 Sta- tus' Habi- tat2 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 0 0 0 0 11 0 0 4 5 Percentage of species introduced to Madagascar 0 2 2 1 0 2 2 0 2 2 0 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) 0 0 810 m 65 43 (66) 61 (94) 43(66) 0 0 1200 m 62 42 (68) 58 (94) 42 (68) 0 0 1500 m 47 30(64) 44(94) 36 (77) 0 0 1875 m 37 23 (62) 34 (92) 24 (65) 0 0 Overall 87 58 (67) 78(90) 52 (69) 0 0 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 days2 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 3 ioo- ! OS i .2; 50- I H Malaza Bealoka k aJJALiBi ih -■- ■ W MP » — — — _ — , 3 to .jo 8 S "S c c: a 2 52 "c S •§ S § * « s § i s .52 co to co co S> C c q> ■C w CO .5 C co ® a oj © to Q Q. 5 6 O to c 5 o o o a 1 -Q CO Co CI c o u 5 CD C S c co 3 CJ CO 3 o .CJ CO a v. ^. * co co <£ •£ S •2 £ .« co c co a *. S ff 9 ■§ S ill a co s O co o i? a a o ** O) CO 2. ■ o rjv .CO Co -O co m co -5 2 « § £ t: Co .Q. CJ Uj 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 463 292 425 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, + N2- 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 birds1 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), N2 = 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 Literature Cited Andersson, M. 1995. Evolution of reversed sex roles, sexual size dimorphism, and mating system in coucals (Centropodidae, Aves). Biological Journal of the Lin- nean Society, 54: 173-181. Andrianarimisa, A. 1994. Note on Tylas eduardi (Ty- las Vanga) at Pare National Montagne d'Ambre. Newsletter Working Group on Birds in the Madagas- car Region, 4: 6. Andrianbololona, A. 1979. L'approvisennement en dnergie de la region de Tulwar. Mdmoire Maitrise, University de Tulear, Toliara, Madagascar. Appert, O. 1966. Beitrag zur Biologie und zur Kenntnis der Verbreitung des Madagaskar Mahnenibisses Lo- photibis cristata (Boddaert). Journal fiir Ornithologie, 107: 315-322. -. 1985. Zur biologie der Mesitornithiformes (Nakas oder "Stelzenrallen") Madagaskars und erste fotografische dokumente von vertretern der ordnung. Ornithologische Beobachter, 82: 31-54. Battistini, R. 1964. Etude gdomorphologique de 1'extreme sud de Madagascar. Editions Cujas, Tou- louse, 340 pp. . 1972. Madagascar relief and main types of landscape, pp. 1-25. In Battistini, R., and G. Richard- Vindard, eds., Biogeography and Ecology in Mada- gascar. W. Junk, The Hague, 765 pp. Bazot, G. 1974. G£ologie de la region Fort Dauphin — Sainte Luce (Sud-Est de Madagascar). Direction des Mines et de 1'Energie, Travaux du Bureau G£olo- gique, Num£ro 142, Antananarivo, Madagascar. Beehler, B. 1982. Ecological structuring of forest bird communities in New Guinea, pp. 837-861. In Gressitt, J. L., ed., Biogeography and Ecology of New Guinea. W. Junk. The Hague. Benson, C. W., J. F. R. Colebrook-Robjent, and A. Williams. 1977. Contribution a l'ornithologie de Madagascar. L'oiseau et la Revue Franchise d'ornithologie, 47: 41-64, 167-191. Berlioz, J. 1946. Oiseaux de la Reunion. Imprimerie Arts Graphiques Modernes, Saint-Denis, Reunion. Besairie, H. 1970. Carte Glologique de Madagascar, 1/500,000. Feuille Ampanihy No. 8. Service Geolo- gique et Centre de lTnstitut Geographique National a Madagascar, Tananarive. Bierregaard, R. O., Jr. 1990. Species composition and trophic organization of the understory bird community in a central American terra firme forest, pp. 2 1 7-236. In Gentry, A. H., ed.. Four Neotropical Forests. Yale University Press, New Haven, Conn. Bluntschli, H. 1932. In den Urwaldern auf Madagas- kar. VI. In sudlichen Bergwald und nach dem Trock- enland Androy. Umschau, 36: 1015-1019. Kadavern Skeletten, Hauten und Balgen etc. Manu- script in American Museum of Natural History, New York. Bourgeat, F 1972. Sols sur socle ancien a Madagascar. Memoires ORSTOM no. 57, Paris. Brenon, P. 1972. The geology of Madagascar, pp. 27- 86. In Battistini, R., and G. Richard- Vindard, eds. Bio- geography and Ecology in Madagascar. W Junk. The Hague. Brisson, M. 1760. Ornithologie ou mdthode contenant la division des oiseaux. Chez C. J.-B. Bauche, Paris. Brown, L. H., E. K. Urban, and K. Newman. 1982. The Birds of Africa, vol. 1. Academic Press, London, xiii + 521 pp. Chaperon, P., J. Danloux, and L. Ferry. 1993. Fleuves et rivieres de Madagascar. Monographic Hy- drologique 10, ORSTOM, Paris. Charles, C. S. 1985. Les Mahafale de 1'Onilahy: Des clans au royaume du XV0 siecle a la conquete colon- iale. These Pantheon-Sorbonne, Paris. Charles-Dominique, P. 1976. Les gommes dans le re- gime alimentaire de Coua cristata a Madagascar. L'oiseau et la Revue Francaise d'ornithologie, 46: 174-178. Collar, N. J., and S. N. Stuart. 1985. Threatened Birds of Africa and Related Islands. International Council for Bird Preservation, Cambridge. Crawford, R. M. M. 1989. Studies in plant survival — Ecological case histories of plant adaptations to ad- versity. Blackwell Scientific Publications, Oxford. Daubeton (le jeune), E. L. 1770-1786. [Planches en- luminees d'histoire naturelle.] In Buffon, G. L. L., Histoire naturelle de oiseaux. L Imprimerie Royale, Paris. Decary, R. 1926. Le Sud-Est de Madagascar. Bulletin Economique de Madagascar, 23: 5-27. . 1927. Contribution a la botanique et a la geo- logie de la region Fort-Dauphin — Andrahomana. Bul- letin de l'Acad^mie Malgache, (nouvelle s6rie) 10: 13-18. Dee, T J. 1986. The Endemic Birds of Madagascar. International Council for Bird Preservation, Cam- bridge. Delacour, J. 1931. Description de neuf oiseaux nou- veaux de Madagascar. L'oiseau et la Revue Francaise d'ornithologie, 1: 473-486. . 1932. Les oiseaux de la mission zoologique Franco-Anglo-Am£ricaine a Madagascar, L'oiseau et la Revue Francaise d'ornithologie, 2: 1-96. Donque, G. 1972. The climatology of Madagascar, pp. 87-144. In Battistini, R.. and G. Richard-Vindard, eds., Biogeography and Ecology in Madagascar. W Junk, The Hague, 765 pp. . 1975. Contribution geographique a l'£tude du climat de Madagascar. Nouvelle Imprimerie des Arts Graphiques, Antananarivo, Madagascar. Dransfield, J., and H. Beentje. 1995. The Palms of Madagascar. Royal Botanic Gardens and International Palm Society, Kew, United Kingdom. GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 117 Du Puy, B., J. P. Abraham, and A. J. Cooke. 1994. Les plantes, pp. 15-29. In Goodman, S. M, and O. Langrand, eds., Inventaire biologique, Foret de Zom- bitse. Recherches pour le Developpement, s6rie Sci- ences biologiques, no. special, Centre d'Information et de Documentation Scientifique et Technique, An- tananarivo, Madagascar. Eboroke, S. 1994. Approche phytosociologique de la parcelle 3 de la RNI d'Andohahela. Travaux de Re- cherches, Laboratoire de Phytobiologie, Universite de Tul6ar, Toliara, Madagascar. Eguchi, K., S. Yamagishi, H. Nagata, M. Nakamura, and V. Randrianasolo. 1992. The mixed-species flocks of forest-living birds in Madagascar, pp. 28-38. In Yamagishi, S., ed., Social Structure of Madagascar Higher Vertebrates in Relation to Their Adaptive Ra- diation. Osaka City University, Osaka, Japan. Farkas, T 1974. On the biology of Monticola imerinus (Hartlaub). Bulletin British Ornithologists' Club, 94: 165-170. Flacourt, E. de. 1658 [reprinted in 1995]. Histoire de la Grande Isle de Madagascar. Edition annotee et pre- sentee par Claude Allibert. INALCO-Karthala, Paris. Foster, M. S. 1975. The overlap of molting and breed- ing in some tropical birds. Condor, 77: 304-314. Glander, K. E., B. Z. Freed, and J. U. Ganzhorn. 1985. Meat eating and predation in captive-born semi-free-ranging Lemur fulvus and caged Lemur ma- caco. Zoo Biology, 4: 361-365. Gibbs, H. L., P. R. Grant, and J. Weiland. 1984. Breeding of Darwin's Finches at an unusually early age in an El Nino year. Auk, 101: 872-874. Godfrey, L. R., W. L. Jungers, K. E. Reed, E. L. Si- mons, and P. S. Chatrath. 1997. Subfossil lemurs: Inferences about past and present primate communi- ties in Madagascar, pp. 218-256. In Goodman, S. M., and B. D. Patterson, eds., Natural Change and Human Impact in Madagascar. Smithsonian Institution Press, Washington, D.C. Goodman, S. M., G. K. Creighton, and C. Raxworthy. 1991. The food habits of the Madagascar Long-eared Owl Asio madagascariensis in southeastern Madagas- car. Bonner zoologische Beitrage, 42: 21-26. Goodman, S. M., and O. Langrand. 1993. Food habits of the Barn Owl {Tyto alba) and the Madagascar Long-eared Owl (Asio madagascariensis) on Mada- gascar: Adaptation to a changing environment. Pro- ceedings of the Pan-African Ornithological Congress, 8: 147-153. Goodman, S. M., O. Langrand, and C. J. Raxworthy. 1993. The food habits of the Madagascar Long-eared Owl (Asio madagascariensis) in two habitats in south- ern Madagascar. Ostrich, 64: 79-85. Goodman, S. M., O. Langrand, and J.-C. Razafima- haimodison. 1994a. Les oiseaux, pp. 73-84. In Good- man, S. M., and O. Langrand, eds., Inventaire biolo- gique, Foret de Zombitse. Recherches pour le Devel- oppement, serie Sciences biologiques, no. special, Centre d'Information et de Documentation Scienti- fique et Technique, Antananarivo, Madagascar. Goodman, S. M., O. Langrand, and B. M. Whitney. 1996. A new genus and species of passerine from the eastern rain forest of Madagascar. Ibis, 138: 153-159. Goodman, S. M., S. O'Connor, and O. Langrand. 1994b. A review of predation on lemurs: Implications for the evolution of social behavior in small, nocturnal primates, pp. 51-66. In Kappeler, P. M., and J. Gan- zhorn, eds., Lemur Social Systems and Their Ecolog- ical Basis. Plenum, New York. Goodman, S. M., and P. Parrillo. In press. A study of the diets of insectivorous Malagasy birds based on stomach contents. Ostrich. Goodman, S. M., and M. Pidgeon. 1991. Madagascar Harrier Hawk Polyboroides radiatus preying on flying fox Pteropus rufus. Ostrich, 62: 215-216. Goodman, S. M., and M. S. Putnam. 1996. The birds of the eastern slopes of the Reserve Naturelle Integra- le d'Andringitra, Madagascar, pp. 171-190. In Good- man, S. M., ed., A Floral and Faunal Inventory of the Eastern Slopes of the Reserve Naturelle Inte- grale d'Andringitra, Madagascar: With Reference to Elevational Variation. Fieldiana: Zoology, new series, 85: 158-170. Goodman, S. M., and D. Rakotondravony. 1996. The Holocene distribution of Hypogeomys (Rodentia: Muridae: Nesomyinae) on Madagascar, pp. 283-293. In Lourenco, W R., ed., Biogeographie de Madagas- car. ORSTOM Editions, Paris. Goodman, S. M., and T S. Schulenberg. 1991. The rediscovery of the Red-tailed Newtonia Newtonia fa- novanae in south-eastern Madagascar with notes on the natural history of the genus Newtonia. Bird Con- servation International, 1: 33-45. Goodman, S. M., D. E. Willard, and P. C. Gonzales. 1995. The birds of Sibuyan Island, Romblon Prov- ince, Philippines, with Particular Reference to Eleva- tional Distribution and Biogeographie Affinities. Fieldiana: Zoology, new series, 82: 1-57. Grandidier, G., 1902. Observations sur les lemuriens disparus de Madagascar. Collections Alluaud, Gaub- ert, Grandidier. Bulletin Museum d'Histoire Naturelle, Paris, 7: 497-505, 587-592. . 1903. Description de V Hypogeomys australis, une nouvelle espece de rongeur sub-fossile de Mada- gascar. Bulletin Museum d'Histoire Naturelle, Paris, 9: 13-15. . 1928. Description de deux nouveaux mammi- feres insectivores de Madagascar. Bulletin Museum National d'Histoire Naturelle, Paris, sen 2, 34: 63-70. Green, G. M., and R. W. Sussman. 1990. Deforestation history of the eastern rainforests of Madagascar from satellite images. Science, 248: 212-215. Griveaud, P. 1960. Une mission de recherche de 1T.R.S.M. au lac Ihotry (S.E. Morombe, Province de Tulear). Le Naturaliste Malgache, 12: 33-41. Gyldenstolpe, N. 1933. A remarkable new flycatcher from Madagascar. Arkiv for Zoologi, 25B(2): 1-3. Halleux, D., and S. M. Goodman. 1994. The redis- covery of the Madagascar Red Owl Tyto soumagnei (Grandidier 1878) in north-eastern Madagascar. Bird Conservation International, 4: 305-311. Hawkins, A. F A., J.-M. Thiollay, and S. M. Good- man. In press. The birds of the Reserve Speciale d'Anjanaharibe-Sud, Madagascar. In Goodman, S. M., ed., A Floral and Faunal Inventory of the Reserve 118 FIELDIANA: ZOOLOGY Speciale d'Anjanaharibe-Sud, Madagascar: With Ref- erence to Elevational Variation. Fieldiana: Zoology. Hoogstraal, H. 1953. Ticks (Ixodoidea) of the Mala- gasy faunal region (excepting the Seychelles). Their origin and host relationships; with descriptions of five new Haemaphysalis species. Bulletin Museum of Comparative Zoology, 111(2): 37-113. Humbert, H. 1935. L'extinction des derniers vestiges de certains types de vegetation autochone a Madagas- car. Archives Museum National d'Histoire Naturelle, Paris, seY 6, 12: 569-587. 1941. Le massif de l'Andohahela et ses d£- pendances (Madagascar, Reserve Naturelle n° XI). Compte rendu sommaire des stances, Socie^ de Bio- geographie. 18: 31-37. Janzen, D. H., M. Ataroff, M. Farinas, S. Reyes, A. Rincon, A. Soler, P. Soriano, and M. Vera. 1976. Changes in the arthropod community along an ele- vational transect in the Venezuelan Andes. Biotropica, 8: 193-203. Jenkins, M. D., ed. 1987. Madagascar: An Environ- mental Profile. International Union for the Conserva- tion of Nature and Natural Resources, Conservation Monitoring Unit, Cambridge and Gland, 374 pp. Jouanin, C, and J.-L. Mougin. 1979. Order Procellar- iiformes, pp. 48-1 18. In Mayr, E., and G. W. Cottrell, eds., Check-list of Birds of the World, 2nd ed., vol. I. Museum of Comparative Zoology, Cambridge, Mass. Kahl, M. P. 1979. Family Scopidae, pp. 244-245. In Mayr, E., and G. W Cottrell, eds.. Check-list of Birds of the World, 2nd ed., vol. 1 . Museum of Comparative Zoology, Cambridge, Mass. Langrand, O. 1990. Guide to the birds of Madagascar. Yale University Press, New Haven, Conn., xi + 364 pp. . 1995. Check List of the Birds of Amber Moun- tain National Park, Madagascar. WWF, Antananarivo, Madagascar. Langrand, O., and O. Appert. 1995. Harlequin Quail Coturnix delegorguei and Common Quail Coturnix coturnix on Madagascar: Occasional migrants or res- ident species? Ostrich, 66: 150-154. Langrand, O., and B.-U. Meyburg. 1984. Birds of prey and owls in Madagascar: Their distribution, sta- tus and conservation. Proceedings of the Second Sym- posium on African Predatory Birds, Natal Bird Club, 1984: 3-13. Langrand, O., and J. C. Sinclair. 1994. Additions and supplements to the Madagascar avifauna. Ostrich, 65: 302-310. Langrand, O., and L. Wilme. 1997. Effects of forest fragmentation on extinction patterns of the endemic avifauna of the Central High Plateau of Madagascar, pp. 280-305. In Goodman, S. M., and B. D. Patterson, eds.. Natural Change and Human Impact in Madagas- car. Smithsonian Institution Press, Washington, D.C. Lavauden, L. 1937. Supplement. In Milne Edwards, A., and A. Grandidier. eds., Histoire physique, natu- relle et politique de Madagascar. Vol. 12, Oiseaux. Soci6t6 d'Editions Geographiques Maritimes et Co- loniales, Paris. Lowry, P. P., II, and D. Faber-Langendoen. 1991. Madagascar Minerals Project, Contract no. MMC- 0093 and MMC-0109. Environmental Impact Assess- ment Study. Part I: Natural Environment. Appendix III: Flora and Vegetation Study. [Copy available in Field Museum of Natural History Library.] Mabberley, D. J. 1989. The Plant-book. Cambridge University Press, Cambridge. MacPhee, R. D. E. 1987. The shrew tenrecs of Mada- gascar: Systematic revision and Holocene distribution of Microgale (Tenrecidae, Insectivora). American Mu- seum of Natural History Novitates, 2889: 1-45. MacPhee, R. D. E., and D. A. Burney. 1991. Dating of modified femora of extinct dwarf Hippopotamus from southern Madagascar: Implications for constrain- ing human colonization and vertebrate extinction events. Journal of Archaeological Science, 18: 695- 706. Marshall, J. T 1978. Systematics of smaller Asian night birds based on voice. Ornithological Mono- graph, no. 25. Mayr, E. 1942. Systematics and the Origin of Species. Columbia University Press, New York. . 1992. A local flora and the biological species concept. American Journal of Botany, 79: 222-238. McCoy, E. D. 1990. The distribution of insects along elevational gradients. Oikos, 58: 313-322. McKitrick, M. C, and R. M. Zink. 1988. Species con- cepts in ornithology. Condor, 90: 1-14. Milne Edwards, A., and A. Grandidier. 1879. His- toire physique, naturelle et politique de Madagascar. Volume XIV Histoire naturelle des oiseaux. Tome I, texte, Paris. Milon, P. 1950. Description d'une sous-espece nouvelle d'oiseau de Madagascar. Bulletin Museum National d'Histoire Naturelle (2e sene), 22: 65-66. . 1952. Notes sur le genre Coua. L'oiseau et la Revue Francaise d'ornithologie, 22: 75-90. Milon, P., J.-J. Petter, and G. Randrianasolo. 1973. Faune de Madagascar. Tome XXXV, Oiseaux. OR- STOM/CNRS, Antananarivo and Paris. Missouri Botanical Garden. 1993. Botanical check- list of Reserve Naturelle Integrate 1 1 (Andohahela). July 1993. Missouri Botanical Garden, St. Louis, Mo. Morat, P. 1973. Les savanes du sud-ouest de Mada- gascar. Memoires ORSTOM no. 68, Paris. Moreau, R. E. 1936. Breeding seasons of birds in East African evergreen forest. Proceedings of the Zoolog- ical Society of London, 1936: 631-653. Nagata, H., E. Urano, K. Eguchi, and S. Yamagishi. 1992. Avifauna of each study sites visited in Mada- gascar, pp. 21-27. In Yamagishi, S., ed., Social Struc- ture of Madagascar Higher Vertebrates in Relation to Their Adaptive Radiation. Osaka City University, Osaka, Japan. Nicoll, M., and Langrand, O. 1989. Madagascar: Re- vue de la conservation et des aires protegees. World Wide Fund for Nature, Gland, Switzerland. Noizet, G. 1953. Carte geologique au 1/100.000 Tran- omaro Marohotro, Service G6ologique, Antananarivo, Madagascar. O'Connor, S. M. 1987. The effect of human impact on vegetation and the consequences to primates in two GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 119 riverine forests, southern Madagascar. Ph.D. thesis, Cambridge University, Cambridge. O'Connor, S., M. Pidgeon, and Z. Randria. 1985. Un programme de conservation pour la Reserve d'Andohahela, pp. 31-36. In Mittermeier, R. A., L. A. Rakotovao, V. Randrianasolo, E. J. Sterling, and D. Devitre, eds. Priorites en matiere de conservation des especes a Madagascar. Occasional Papers of the IUCN Species Survival Commission, Gland, Switzerland. Office National de l'Environnement. 1994. Rapport sur l'6tat de l'environnement a Madagascar. Pro- gramme des Nations Unies pour le Developpement Banque Mondiale, Antananarivo, 208 pp. Olson, D. M. 1994. The distribution of leaf litter in- vertebrates along a Neotropical altitudinal gradient. Journal of Tropical Ecology, 10: 129-150. Paulian, R., C. Blanc, J.-L. Guillaumet, J.-M. Betsch, P. Griveaud, and A. PEYRifiRAS. 1973. Etude des eco- systemes montagnards dans la region malgache. II. Les chames Anosyennes. Geomorphologie, climato- logie et groupements vegetaux. (Campagne RCP 225, 1971-1972). Bulletin Museum National d'Histoire Naturelle, Ecologie generate, third series, 118: 1-40. Perrier de la Bathie, H. 1936. Biogeographie des Plantes de Madagascar. Societe d'Edition Geograph- iques, Maritimes et Coloniales, Paris. Peters, D. S. 1996. Hypositta perdita n. sp., eine neue Vogelart aus Madagaskar. Senckenbergiana Biologica, 76: 7-14. Peyrot, B. 1980. La vie rurale en pays Antanosy. Mad- agascar, Revue de Geographie, 37: 111-138. Poulsen, B. O. 1994. Mist-netting as a census method for determining species richness and abundances in an Andean cloud forest bird community. Gerfaut, 84: 39-49. Prigogine, A. 1980. The altitudinal distribution of the avifauna in the Itombwe Forest (Zaire). Proceedings of the Pan- African Ornithological Congress, 4: 169- 184. Putnam, M. S. 1996. Aspects of the breeding biology of Pollen's Vanga (Xenopirostris polleni) in south- eastern Madagascar. Auk, 113: 233-236. Rakotoarisoa, J. A. 1994. Le peuplement de l'Anosy, les rapports de l'homme et son environnement, dans 1' extreme sud-est de Madagascar. These soutenue a 1TNALCO, Paris. 1997. A cultural history of Madagascar: Evo- Zoologique Franco-Anglo-Americaine a Madagascar. Bulletin American Museum of Natural History, 72: 143-499. 1951. The nests and eggs of Mesoenas uni- lution and interpretation of the archaeological evi- dence, pp. 331-341. In Goodman, S. M., and B. D. Patterson, eds., Natural Change and Human Impact in Madagascar. Smithsonian Institution Press, Washing- ton, D.C. Rakotondranony, L. G. 1977. A propos de Berenty inventaire de l'avifaune notice sur les vertebres non primates. Memoire de fin d'etudes, Universite de Madagascar, Etablissement d'Enseignement Superieur des Sciences Agronomiques, Departement Eaux et Forets. [Copy available in the Field Museum of Nat- ural History Library.] Rand, A. L. 1933. Testicular asymmetry in the Mada- gascar Coucal. Auk, 50: 219-220. . 1936. The distribution and habits of Madagas- car birds. A summary of the field notes of the Mission 120 color of Madagascar. Auk, 68: 23-26. 1960. Family Vangidae, pp. 365-369. In Mayr, E., and J. C. Greenway, Jr., eds., Check-list of Birds of the World, vol. 9. Museum of Comparative Zool- ogy, Cambridge, Mass. Ratsirarson, J. 1996. Importance de la recherche au sein du projet Beza Mahafaly. Akon'ny Ala, 18: 12- 25. Ratsirarson, J., J. A. Silander, Jr., and A. F. Richard. 1996. Conservation and management of a threatened Madagascar palm species, Neodypsis decaryi, Jumelle. Conservation Biology, 10: 40-52. Ratsivalaka-Randriamanga, S. 1985. Recherches sur le climat de Tolagnaro (ex Fort-Dauphin) (Extreme Sud de Madagascar). Madagascar, Revue de Geogra- phic 46: 47-67. . 1987. Climat et vegetation de la region de Fort-Dauphin. Recherches pour le Developpement, serie Sciences de 1' Homme et de la Societe, 3(premier semestre): 51-64. Razanabahiny, V. 1995. Le Dina (Convention entre Membres de Communautes Villageoises) son oppor- tunity ou non dans la conservation de la nature. Cas de la Reserve Naturelle Integrate d'Andohahela — To- lagnaro. Memoire C.A.P.E.N., Ecole Normale Super- ieure, Universite d' Antananarivo, Antananarivo, Mad- agascar. Remsen, J. V., Jr. 1994. Use and misuse of bird lists in community ecology and conservation. Auk, 111: 225- 227. Remsen, J. V., Jr., and D. A. Good. 1996. Misuse of data from mist-net captures to assess relative abun- dance in bird populations. Auk, 113: 381-398. Remsen, J. V., Jr., and T. A. Parker III. 1983. Contri- bution of river-created habitats to bird species richness in Amazonia. Biotropica, 15: 223-231. Richard, A. F, and R. E. Dewar. 1991. Lemur ecol- ogy. Annual Review of Ecology and Systematics, 22: 145-175. Roche, H. de la, and J. Marchal. 1955-1956. Carte geologique au 1/200.000 Manantenina Fort- Dauphin, Service Geologique, Antananarivo, Madagascar. Safford, R. J., and J. W Duckworth, eds. 1990. A Wildlife Survey of Marojejy Nature Reserve, Mada- gascar. ICBP Study Report No. 40. International Council for Bird Preservation, Cambridge. Salomon, J.-N. 1993. La deforestation a Madagascar une dynamique inquietante. Publication presentee a l'Universite d'Hiver — Aupelf-Uref, mai 1993. Univ- ersite d' Antananarivo. Probleme de l'environnement en milieu tropical dans les ties de l'ouest de l'Ocean Indien. Salomonsen, F. 1933a. Les gobe-mouches de paradis de la region malgache. L'oiseau et la Revue Francaise d'Ornitholgie, 3: 603-614. . 1933b. Remarks upon the Madagascar Paradise Flycatchers. Bulletin of the British Ornithologists' Club, 53: 119-124. FIELDIANA: ZOOLOGY . 1934. Revision of the Madagascar Timaliine birds. Annals and Magazine of Natural History, series 10, 14: 60-79. Salvan, J. 1970. Remarques sur Involution de l'avifaune malgache depuis 1945. Alauda, 38: 191- 203. Sauther, M. L. 1989. Antipredator behavior in troops of free-ranging Lemur catta at Beza Mahafaly Special Reserve, Madagascar. International Journal of Prima- tology, 10: 595-606. SCHULENBERG, T. S., S. M. GOODMAN, AND J.-C. RAZA- fimahaimodison. 1993. Genetic variation in two sub- species of Nesillas typica (Sylviinae) in south-east Madagascar. Proceedings of the Pan-African Ornitho- logical Congress, 8: 173-177. Smrihai, H., I. Sinclair, and P. R. Colston. 1995. A new species of Puffinus shearwater from the western Indian Ocean. Bulletin of the British Ornithologists' Club, 115: 75-87. Snow, D. W., ed. 1978. An Atlas of Speciation in Af- rican Non-passerine birds. British Museum (Natural History), London, vii + 390 pp. Snow, D. W., and B. K. Snow. 1964. Breeding seasons and annual cycles of Trinidad land-birds. Zoologica, 49: 1-39. Storer, R. W. 1966. Sexual dimorphism and food hab- its in three North American accipiters. Auk, 83: 423- 436. Stresemann, E. 1952. On the birds collected by Pierre Poivre in Canton, Manila, India and Madagascar (1751-1756). Ibis, 94: 499-523. Terborgh, J. 1971. Distribution and environmental gra- dients: Theory and a preliminary interpretation of dis- tributional patterns in the avifauna of the Cordillera Vilcabamba, Peru. Ecology, 52: 23-40. . 1977. Bird species diversity on an Andean ele- vational gradient. Ecology, 58: 1007-1019. Thorstrom, R., and R. Watson. 1994. Avian Inven- tory of Masoala Peninsula. Madagascar. The Peregrine Fund, Boise. Id. UlLENBERG, G., H. HOOGSTRAAL, AND J. M. KLEIN. 1979. Les tiques (Ixodoidea) de Madagascar et leur role vec- teur. Institut Pasteur de Madagascar, Numero Special. Vernon, C. J. 1971. Notes on the biology of the Black Coucal. Ostrich, 42: 242-258. Walker, A. 1967. Locomotor adaptations in recent and subfossil Madagascan lemurs. Ph.D. thesis. University of London. London. Watson, R. T, and S. Strzalkowska. 1992. Project Masoala: Les trois prochaines annees, pp. 87-106. In Watson, R. T, ed.. Le projet de conservation de l'aigle pecheur de Madagascar et des zones humides et le projet Masoala: Utilisation des rapaces et d'autres fau- nes pour la gestion et revaluation de la conservation. The Peregrine Fund. Boise, Id. White, F. 1983. The Vegetation of Africa. UNESCO, Paris. Wilkinson, R. 1990. Notes on the breeding and behav- iour of Greater Vasa Parrots Coracopsis vasa at Ches- ter Zoo. Avicultural Magazine. 96: 1 15-122. Wilkinson, R., and T R. Birkhead. 1995. Copulation behaviour in Vasa parrots Coracopsis vasa and C. ni- gra. Ibis, 137: 117-119. Wright, H. T, and J. A. Rakotoarisoa. 1997. Cultural transformations and their impact on the environments of Madagascar, pp. 309-330. In Goodman, S. M., and B. D. Patterson, eds.. Natural Change and Human Im- pact in Madagascar. Smithsonian Institution Press. Washington. D.C. Wright, H. T, J. A. Rakotoarisoa, G. Heurtebize, and P. Verin. 1993. The evolution of settlement systems in the Efaho River Valley, Anosy: A preliminary re- port on archaeological reconnaissances of 1983-1986. Indo-Pacific Prehistory Bulletin, 12: 2-20. Yamagishi, S., and K. Eguchi. 1996. Comparative for- aging ecology of Madagascar vangids (Vangidae). Ibis, 138: 283-290. Yamagishi, S., E. Urano, N. Rakotoarison, H. Ran- driamahazo, and V Randrianasolo. 1992. A pos- sible significance of dichromatism in male Madagas- car Paradise Flycathers (Terpsiphone mutata), pp. 58- 61. In Yamagishi, S., ed.. Social Structure of Mada- gascar Higher Vertebrates in Relation to Their Adap- tive Radiation. Osaka City University, Osaka, Japan. GOODMAN ET AL.: BIRDS OF SOUTHEASTERN MADAGASCAR 121 Appendix 1. Gazetteer of localities mentioned in the text. Eastern latitude Southern longitude Elevation Locality 0 ' 0 ' (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 0 ' 0 ' (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 /' 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 Fowj 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 M t'il K .'i:k1 ;H I »( o- 249ft —