Biogéographie de , 1996 :349-362

COMPOSITION AND CHARACTERISTICSOF BlRD COMMUNITIES IN MADAGASCAR

Lucienne WILME

Missouri Botanical Garden - Madagascar Research andConservation Program, BP 3391, Antananarivo 101, MADAGASCAR

ABSTR4CT.- Currentknowledge of the taxonomicand geographic composition of Madagascar's resident avifauna is reviewed, with particular emphasis on forest . Distribution patterns in the avifauna do not match those of the botanical communities, and especially the domains of floristic endemism in the Eastern Region. The toleranceof the endemic forest avifauna to habitat degradationis proportional to its degree of taxonomic endemism. The21 belonging tothe two endemic families and the two endemic subfamilies are clearly the most sensitive to forest degradation. The members of thesehigher taxonomic groups could serve as usefulbiological indicators of Madagascar'sforest habitats.

KEY W0RDS.- Madagascar, Birds, Endemjsm, Ecological niche, Biological indicator

RÉSUMÉ.- Ce travail présente un résumé dela connaissance actuelle dela composition taxinomique et géographique de l'avifaune résidente de Madagascar et plus particulièrement de l'avifaune forestière. Les schémas de distribution de l'avifaune ne se superposent pas à ceux des communautés botaniques et notammenten matière de description des domaines d'endémisme floristiques de la Région Est de Madagascar. En considérantl'avifaune endémique forestière de Madagascar, la tolérance à la dégradation des forêts est proportionnelleau degré d'endémisme taxinomique, et il apparaît que les 21 espècesappartenant aux deux familles et deux sous-familles endémiques à Madagascarsont certainement les plus sensiblesà la dégradation des forêts. L'ensemble des éléments appartenant à ces taxons supérieurs devraient constituer les indicateurs biologiques des milieux forestiers de Madagascar.

MOTS-CLES.- Madagascar, Oiseaux, Endémisme, Niche écologique, Indicateur biologique

INTRODUCTION

On the basis of floristic and ecological data, Madagascar can be divided into two majorphytogeographic regions, the East MalagasyRegion and the WestMalagasy Region (HWERT, 1955). According to WHITE (1983), the East Malagasy Region is divided into four domains: Eastern (escarpment and coastal zone), Central (eastern part of central highlands), High Mountain, and Sambirano; and the West Malagasy Region into the Western and Southern Domains (Table 1, Fig. 1). Details on sites within the existing protected areas system and those of biological interest are reviewed in NICOLL and LANGRAND(1989). A total of 204 breedingspecies of birds are recognized in Madagascar. This includes the 201 species listed in LANGRAND(1990), minus the Snail-eating Coua

In: W.R. LOURENÇO (éa.) Editions de I'ORSTOM, Paris 350 L. WJLME delalandei, now regarded as an extinctspecies (GOODMAN, 1993a),plus four new additions to the list: the CommonQuail Coturnixcoturnix and the HarlequinQuail Coturnixdelegorguei now consideredas nesting species (LANGRAND& APPERT,in press.), a new Sylviidae being described(GOODMAN et al., in press.), and NesiZlas typica lantzii, which is being elevatedto the rank of species (SCHULENBERGet al., 1993). Three of these 204 species have been introduced to the island: Common Myna Acridotheres tristis (at the end of last century), Common WaxbillEstrilh astriZd in 1983, and House Sparrow Passer domesticus in 1984 (LANGRAND,1990). A total of 106 (53%) of the 201 non-introduced species are endemic to Madagascar and another 25 species (just over 12%) havea distribution limited to awider region including Madagascar and the neighboring islands (Comoros, Mascarenes, andthe Seychelles) (LANGRAND,1990). In view of the declining forests of Madagascar (JENIUNS, 1987; GREEN& SUSSMAN,1990; NELSON& HORNING, 1993a, 1993b) an understanding of the resident avifauna's geographic distribution, niche occupancy and tolerance to habitat disturbance might help to identify conservation priorities andto predict responses of the avifauna to human disturbance. The high degree of endemism among this rather mobile group of not only justifies specialattention in terms of conservation efforts, but also offers an ideal set of species to test models in community ecology.

Montagne bm ICI

Western Domain

I 11 Andohahela (humid)

Fig. 1. Location of study siteswithin phytogeographicdivisions; Eastern Region; Southern Domain; Western Domain; Abbreviation as in Table II. C0M"ITES IN WAGASCAR 351

TAXONOMIC AND GEOGWHIC DISTRIBUTION

Of the 201non-introduced breeding species66 are Passeriformes, whichare under- represented outside forest habitats.Within forests theycomprise a high percentage (90.5%) ofthe endemicpasserine species in Madagascar (Fig. 2). Within each of Madagascar's regions and domains breeding species are mainly found in forest habitats emphasizing the predominant role of this type of habitat for the Malagasy avifauna. This role is highlighted bythe fact that 85 (80%) of al1 endemic species relyon forest habitats. Considering only the 114 forest birds species, this meansthat three out of four species in Malagasy forests are endemic (Fig. 2). For this reason, the present analysis is restricted to species occuring in natural forests,though other habitats certainly also deserve special attention (LANG- & WL~,1993).

48 rn 44 .-a, O a, Q u) rc O 55 d E 3 IO 7 8

forests open aquatic marineaquatic open forests Main habitats

endemicpasserinenon endemic passerine non endemic non passerine0 non endemic passerine Fig. 2. Composition of Madagascar nesting bird species according to main habitat (numbers are total numberof species per habitat).

The classification summarized in Table II shows the distribution of forest birds according to the phytogeographic regions and domains.The Eastern Region as a whole contains more species and more regional endemism thanthe Western Region; 99 species, with 36 endemic in the East, versus 77 and 15 regional endemics in the West. This pattern holds for al1 forest bird taxa, but is more marked for passerines. Neither the 352 L. WILME

Eastern, Sambirano, nor High Mountains Domains hold any local endemics. Only the Central Domain contains local endemics (five species). In contrast both domains of the Western Region have local endemic species(two in the Western Domain and nine inthe Southern Domain). To understandsimilarities in speciescomposition and assess species turnover, Jaccard's index was calculated separately for endemic, non-endemic, non-passerine and passerine species for each of the 22 sites studied (Table III). This index for comparison between two sites is calculated as

Nc JI= where Nc is the number of species commonat both sites; andNA and NB are the numbers of species present at site A and site B respectively (MUELLER-DOIVBOIS& ELLENBERG, 1974). The sites used in this calculationare shown in Figure 1 along with the domain in which they occur. Species listsare based on atleast 30 days of intensive surveysper site. Cluster analysis with Jaccard's indices of similarity between sitesdoes not produce geographically distinct clusters when applied to non-endemic species. This means that among non-endemic birds,there is no distinct set of species inhabiting dry forests versus those predominantly in wet forest areas. Non-endemic species have broad distributions and low turnover rates, indicating more generalizsd habit requirements. Applyingcluster analysis to similarityindices of Madagascar endemic species reveals three distinctclusters. The geographicextent and characteristics of species similarities within and betweenthese clusters are shown in figure 3. The endemic avifauna differs little between sites of the eastern rainforest. The humid forest parce1 of Andohahela (Parcel 1) in the extreme south holds the same island endemicbird species as Marojejy in the northeast.This homogeneity in species composition ends abruptly along the western watershed of Andohahela. Here species turnover is almost complete between the humid Eastern Region and the dry Southern Domain,which is itself homogeneous in terms of endemicspecies in general and passerines in particular(two non-passerines species have a limited range inthe Southern Domain, Monias benschii, Uratelornis chimaera,and Coua verreauxi does not occur in the eastern part of the Domain). The avifauna of Isalo, with its locally endemic passerine (Pseudocossyphzrs bensoni)is more closely relatedto that of the south and the West than to that of the Central Domain, contrary to expectations on phytogeographicgrounds. The species assemblage, especially for passerines, changes between sites of the SouthernDomain and the WesternDomain, as represented by Zombitse, Kirindy, Bemaraha, Ankarafantsika and Ankarana. Montagne d'Ambre and Manongarivo do not cluster with the other sites: Montagne d'Ambre is an isolated island of raidorest within the Western Domainand its avifaunais depauperate; Manongarivo belongs to the SambiranoDomain and its birdcomposition shows affinities with both eastern and western sites. Here we find eastern speciessuch as Cozra caerulea, C. serriana and Phyllastrephus zosterops together with western species such as Falculea palliata and Philepitta schlegeli. Even though the species list for Manongarivo is not complete, the site's geographic position seemsto overide phytogeographic effects. BIRD COMMUNITIES IN MADAGASCAR 353

DISTRIBUTION ACCORDING TO NICHES

Species were assigned to niches according to their main food items, and their feeding and nesting habits (TableII). On the basis of these niche variables, the 114 forest bird species are distributed among 28 different niches. To facilitate comparisons, each of the 28 niches was assigned to one of the 16 categories listed in Table IV. The absolute numbers of birds species increase with increasingstructural and floristic diversity of the forest, f?om the drier western forests to the wet forests of the center and east. However the number of guilds and the relative number of species in each guild remain similar in both phytogeographicregions. Insectivores, followed bysmall predators, are the dominant groups throughout the island. The main difference betweenthe Eastern and the Western Region is the higher representation of insectivores inthe east. Whereas the east holds 39 of the 45 strictly insectivorous forest species, the West contains only 23 of 45. Though forest structure is correlated with bird species diversity,it alone is insufficient to explain the greater species richness in the east, which is mainly due to an increase in arborealspecies. This isexemplified by the highernumber of strictly arboreal insectivorous species inthe Central Domain with forest a height of up to 25 m, compared to thelower number of species in this guild inforests of approximately the same height in the Eastern Domain andthe Sambirano.

Fig. 3. Species similarities within andbetween separate units of the endemic avifauna as identified by cluster analysis. Values are means of Jaccard's indices. The upper and lower value of each pair of numbers refer to endemic non passerine and endemic passerine respectively. 354 L. WILME

Within the Eastern Region, the Eastern Domainis somewhat depauperate in insectivores (Table IV). Seven of the 10 species common to the Eastern region, but not present in the Eastern Domain itself, are insectivores. The Sambirano links and combines nicheelements belonging to the Eastern or to the WesternRegion. Both the High Mountain Domain and the Sambirano hold a reduced set of species that are widespread across the island,and contain no localendemic bird species. Among the forest bird species, taxonomical endemism at the species level is particularly high in the guild of exclusivelyarboreal insectivorous passerines. The geographic distribution of these groups of endemicspecies is uneven. Whereas insectivores (such as the PhyZZastrephus) and predators reached higher species numbers thein east, omnivores and (( vegetarian >) species have aboutthe same number of species in the east and in the West, respectively 16 (16%) and 10 (10%) for the east, and 15 (19%) and 10 (13%) for the West. This difference mightbe linked to the pronounced dry seasonof the West, favoring species whichare able to feed on agreater variety of food items.

TOLERANCETO FOREST DISTTJRBANCE

Four categories are recognized to describe the quality of forest habitats: undisturbed forest, slightlydisturbed forest, secondary growth andanthropogenic wooded grassland. Forest species are defined as those occurring in undisturbed forest, although they may also exist in other forest types (Table II). Out of the 114 species occurring in undisturbed forest, 22 (19%) rely exclusively on this habitat, whereas 38 species (33%) also occur in slightly disturbed forest,and another .54(47%) also occur in secondary growth or anthropogenic grassland.

92 spp. r_! 54 spp. undisturbedslightlydisturbed secondary growthsavanna Forest quality

Spp. belongingto an endemic family Spp. belonging to an endemicsubfamily nSpp. belonging to an endemic genera 13 Other endemic spp. .!.:y;, Non endemic spp. with endemic sspp. ,y#% Other non endemic SPP. D,,,/,;+ Figure 4. Taxonomic level of endemism and toleranceto forest disturbanceof Madagascar forest bird species (numbers are total number of species per forest habitat). BIRD COMMUNITIES IN MADAGASCAR 355

Members of Madagascar’s endemic bird familiesand subfamilies are not tolerant of forest disturbance.Of the 32 endemic forest genera, 27 do not occur insecondary growth or anthropogenicgrassland. Onlyeight species (out of 59, belonging to 5 endemic genera occur in secondary habitats. At the sub-family level, only two species of Coua are tolerant of some forest disturbance. The eight species belonging to the two endemic families occur only in undisturbed and slightly disturbedforest (Table V, qg. 4). Studied forests that are not isolated, that cover at leastseveral hundred km ’ with altitudes startingat least at 800 meters above sea levelhold at least 60% of the species in endemic genera withintheir respective phytogeographic region. The information at hand is not sufficient to reveal causal factors in the patterns of species distributions across Madagascar. Current fkagmentation .and previous corridors between the east and the West (e.g. Sambirano) confound the analysis with cases such as the presence of an << eastern >) rainforest species of (Caniraklus kioloides) in the deciduousdry forests of Bemaraha orthe westernspecies of (Mesitornis variegata) in Ambatovaky rainforest in the east. The Malagasy avifaunadoes not reflect the phytogeographic patterns seen among the botanical communities, especially in the Eastern Region.

CONCLUSION

Despite some uncertainties and ambiguities, the present analysis of the Malagasy avifaunare-emphasizes the pending threat to the island’shigher endemic bird taxa, especially the endemic families and sub-families. Species belonging to endemic families (Mesitornithidae, Brachypteraciidae) already have a patchy distribution, andthe modern distributionof three of eightspecies in these families (Monias benschi, Mesitornzs variegata, Uratelornis chimaera) ishighly restricted. The endemic sub-families (Couinae,Phillepittinae) are widespread,although the latter is relativelysensitive to forest disturbance and fragmentation. The Couinae contain a single genus (Coua) with nine extant species that are broadly distributed in different typesof forest; they feed on a variety of food and occupy different forest strata. But the largest species within this sub- family (Coua delalandei, Couaprimavea and Coua berthae) are al1 extinct (GOODMAN, 1993a, 1993b). Undisturbed forests with an area of several thousand ha hold at least three species of Coua belonging to at least two differentecological niches. The distribution pattern of the remaining species is not simple; for example the southern endemic Coua verreauxi has a narrow distribution, while the eastern species C. serriana and eastern subspecies of C. cristata are more common at lower elevation (below 500 m). These higher endemic taxa, with eight genera, 21 species, and six sub-species, may serve as biological indicators or keyelements to defineconservation priorities. The ecological niches utilized by these two families and two subfamilies include insectivores, omnivores,vegetarians and small predators aswell as terrestrial, arboreal, and terrestriaVarborealspecies when considering feeding and nesting habits. Using these speciesand subspecies for conservationpurposes may providea usefùl predictive indicatorof forest conditionsand thus offersa tool for improvedmanagement and preservation of Malagasy naturalforest ecosystems. 356 L. mm

ACKNOWLEDGEMENTS

1 am very gratefbl to Jorg Ganzhorn Who helped analyze the data and improved significantly previous versions of the manuscript. 1 would also like to thank Steve Goodman and Dominique Halleuxfor usefùl suggestions, andPete Lowry for assistance with improvingthe final manuscript.

REFERENCES

GOODMAN, S.M., 1993a. A reconnaissance of Ile Sainte Marie, Madagascar: the statusof the forest, avifauna, lemurs and fruit bats. Biol. Conservation,65: 205-212. GOODMAN,S.M., 1993b. Identification of bird subfossils from cave surface deposits at Anjohibe, Madagascar with a description of a new (Cuculidae: Couinae). Proc. Biol. Soc. Wash., 106(1): 24-33. GOODMAN, S.M., O. LANGRAND & B. WHITNEY. (In press). A new genus and species of passerine from the eastern rainforestof Madagascar. Ibis. GREEN,G.M. & R.W.SUSSMAN, 1990. Deforestation history of theeastern rain forests of Madagascar from satellite images. Science, 248: 212-215. HUMBERT, H., 1955. Les territoires phytogéographiques de Madagascar. In: Les divisions écologique du monde. pp. 195-204. ENKINS, M.D.,1987. Madagascar: An environmental profile. Gland: International Union for the Conservtion of Nature and Natural Resources. LANGRAND, O., 1990. Guide to the birdsof Madagascar. New Haven: Yale University Press. LANGRAND, O. & O. APPERT (In press). Harlequin delegorguei and Common Quail Coturnix coturnix on Madagascar: occasional migrants or resident species? Ostrich. LANGRAND, O. & L. WILME, 1993.Protection des zones humides et conservationdes espèces d'oiseaux endémiques de Madagascar. Proc.VI11 Pan-Afr. Orn. Congr.: 201-208. MUELLER-DOMBOIS, D. & H. ELLEMERG, 1974. Aims and methods of vegetation ecology. New York, John Wiley & Sons, NELSON,R. & N.HORNING, 1993a. Cover foresthon-forest classification of Madagascar fiom AVHRR data. Int. J. Remote Sensing, 14 (8): 1445-1446. NELSON, R. & N. HORNING, 1993b. Am-LACestimates of forest area in Madagascar, 1990. Int. J. Remote Sensing, 14 (8): 1463-1475. NICOLL, M.E. & LANGRAND, O., 1989. Madagascar: Revue dela conservation et des aires protégées. WWF, Gland, Switzerland. xvii 374 pp. + .. PHILLIPSON, P.B., 1994. Indian Ocean: CPD site 101 MADAGASCAR In: S.D. Davis, V.H. Heywood & A.C. Hamilton (eds.) Plant Diversity. A guideand strategy for their conservation. Volume 1. Europe, Afiica, South West Asia andthe Middle East. pp. 271-281. World Wide Fund for Nature (WWF) and The World Conservation Union (IUCN). 358 L.wILME ,

Distribution: level of endemism Ecological niche: w s c eW hmsa E Food Feeding Ne geographic 1 taxonomic rance habit st NON-PASSERINE SPECIES Lophotibîs cristata 11111111 Mad. genera 2 IGV Aviceda madagascariensis lllll Mad. species 3 IVAr R Machaeamphusalcinus 1 111 II1 subspecies 3 IV R Ar Eutriorchis astur 11 1 E genera O V Ar R Polybomides radiatus 11111111 Mad. species 2 V Ar R Accipiter henstii 1 111 1 Mad. species R Ar Accipitermadagascanensis 111111 1 Mad. species Ar R Accipiter francesii 11111111 Ar R Bute0 brachypterus 11111111 Mad. species Ar R Falco zoniventris 111111 1 Mad. species 3 IVAr R Falco peregnnus 11111 11 3 V RF Numida meleagris 111111 1 T 3T Veg Mesifomis variegata 1 11 1 Mad. family O I Veg T Ar Mesitomis unicolor 11 1 E family OAr T I Veg Monias benschi 11 S family OAr T I Veg Tumix nigricollis lllll 1 Mad. species 3 lVeg T T CaniraNus kioloides 1 1111 1 Mad. species 1 I T Ar Samthrura inSulans 11 11 E species 3 lVegT T Ptemcles personatus Ill W speciesT 3T Veg Streptopelia pictufafa 111111 1 subspecies 3T Veg Ar Tremn australis 111111 1 subspecies 3 Ar Veg Ar Alectmenas madagascariensis lllll E Ar 2 Ar Veg Coracopsis vasa 111111 1 subspecies 3 Ar Veg Ar Coracopsis nigra 11111111 subspecies 3 Ar Veg Ar Agapomis cana llllll 1 Mad. speciesAr 3 Ar Veg Cuculus audeberti 11 1 subspecies 1 PI Ar Cuculus mchii 111111 1 Mad. species 3 PI Ar Coua gigas Ill W subfamily 1 I Veg T Ar Coua coquereli II 11Mad. subfamily 1 T 1 Veg Ar Coua seniana 111 1 E subfamily 1 I Veg T Ar Coua reynaudii 11111 E subfamily 1 lVegAr T Coua cursor 11 S subfamily 1 I T Ar Coua Nticeps 111 W subfamily 1 I Veg T Ar Coua crislafa 111111 1 Mad. subfamily 2 IVegGVAr Ar Coua vernaux; 11 S subfamily 1 Ar ArI Veg Coua caerulea 1 111111 Mad. subfamily 2 IVegV Ar Ar Centmpus foulou 111111 1 subspecies 3 IVAr Ar THOsoumagnei 11 1 E species RO V Ar OtUS Nti1U.S 111111 1 subspecies 3 I R Ar Ninox supercilaris 11111 1 Mad. species 2 I RT Asio madagascariensis 111111 1 Mad. species 3 V R Ar Caprimulgus madagascanensis 111111 1 subspecies 3 I Ar T Caprimulgus enamtus 1 Ill 1 Mad. species 1 1 Ar T Zoonavena grandidien 111111 1 subspecies 1 I Ar Ae BlRD COMMUNITIES IN MADAGASCAR357

SCHULENBERG,T.S., S.M. GOODMAN & J.-C.RAZAFIMAHATMODISON, 1993. Genetic variation in two subspecies of Nesillas fypica (Sylviinae) in south-east Madagascar. Proc.VZII Pan-Afr. Om. Congr.: 173-177. WTE, F., 1983.The vegetation of Africa.descriptiveA memoir accompanyto the UNESCO/AETFAT/UNSOvegetation map of Africa.Natural Resources Research. Paris: UNESCO.

Table 1. Biogeographic regions and domainsin Madagascar

RegionsDomainsand Elevation Rainfall season Dry Character diversitySpecies endemismcanopy

Eastern Malagasy Region

EasternDomain0-800m m25-30 >high evergreen 2000high veryno mm CentralDomain300m800-1 > 1500 mm evergreenhigh nohigh 20-25 m 1300-2000 m m 10-12 High Mountain Domain > 2000m substantialevergreen no low veryhigh 6 m Sambirano Domain 0-1876m > 2000 mmevergreen no high high 25-30 m Western Malagasy Region

Western Domain 0-800m 500-2000mm < 7 monthsdeciduous medium very high 12-15 m Southern Domain 0-400m 350-750mm 7 monthsdeciduous medium very high 3-6 m adapted from Phillipson (1994) BIRD C0M"TIES IN MADAGASCAR 359

~~~ ~ Distribution: level ofendemism Ecological niche: w s W c e hmsa E Nest Food geographic 1 taxonomie z:i Apus melba 11111111 lsubsoecies 3 I Ae C Apus barbatus 11111111 subspecies 3 I Ae cl lspidina madagascariensis 111111 1 Mad. species 1 IV Ar T Mempssuperciliosus . 1 1 1 1 1 1 1 3 I Ae T Eufysfomusglaucurus 111111 1 subspecies 3 I Ae Ar Brachypteracias leptosomus 11 1 family 1 IGV T T Brachypteracias squamiger 11 1 family 1 IGV T T Atelomis piftoides 11111 family 1 IV Atelomis cmssleyi 11 1 family O I Uratelomis chimaera II family TT Leptosomus discolor 111111 1 Ar Ar Upupa epops 11111 1 T Ar PASSERINE SPECIES subspecies Philepitta castanea 11111 E subfamily Ar Ar Philepitta schlegeli 1111 Mad. subfamily Ar Ar Neodrepanis coruscans 1111 E subfamily Ar Ar Neodrepanis hypoxantha 1 11 E subfamily Ar Ar Phedina bohonica 11111111 subspecies Ae c Coracina cinerea 11111111 subspecies Ar Ar Phyllastrephusmadagascariensis 1 1 1 1 1 1 1 Mad. species Ar Ar Phyllastrephus zostemps 111 1 E species 1 I Ar Ar Phyllastrephus apperti 11 W species O 1 Ar Ar Phyllastrephus tenebmsus 11 1 E species O I Ar Ar Phyllatrephus cinereiceps 1 1 C species O I Ar Ar Hypsipetesmadagascariensis 1 1 1 1 1 1 1 1 subspecies 3 tVeg Ar Ar Copsychus albospecularis 111111 1 Mad. species 2 I Ar Ar Pseudocossyphus sharpei 1111 E genera 1 lVeg Ar Ar Pseudocossyphus imerinus 11 S genera 1 Veg Ar Ar Pseudocossyphusbensoni 1 1 C genera 2 lVeg Ar Ar Nesillas typica 11111111 subspecies 3 I Ar Ar Nesillas lanfrii 11 S species 3 I Ar Ar Thamnomis chlompetoides 11 S genera 1 I Ar Ar Dmmaeocercus brunneus 1 1 C genera Ar Ar Randia pseudozostemps 11 1 E genera :11 iI Ar Ar Sp. nov. 11 1 E genera Ar Ar Newtonia amphichma 11 1 E genera Ar Ar Newtonia brunneicauda 11111111 Mad. genem OI1 1 Ar Ar Newtonia archboldi 11 S genera Ar Ar Newtonia fanovanae 11 1 E genera Ar Ar Neomixis tenella 111111 1 Mad. genera Ar Ar Neomixis viridis 1111 E genera O3 jI Ar Ar Neomixis striatigula :l 111111 1 Mad. genera 1 I Ar Ar Hartertulaflavovirdis 1 1 C genera O I Ar Ar Pseudobias wardi 1111 E genera O I Ar Ar Teipsiphone mutata 11111111 subspecies 3 I Ar Ar Oxylabes madagascariensis 11 1 E genera 1 I Ar Ar Cmssleyia xanthophrys 1 1 C genera 1 I T T Mystacomis cmssleyi 11 1 E genera 1 I T Ar Nectarinia souimanga 11111111 subspecies 3 l Veg Ar Ar Nectarinia notata 11111111 subspecies 3 lVeg Ar Ar Zostemps maderaspatana 11111111 3 lVeg Ar Ar Calicalicusmadagascariensis 1 1 1 1 1 1 1 Mad. genera 1 IV Ar Ar Schetba Nfa 1 111 1 Mad. genera O IV Ar Ar Vanga cutvimstris 111111 1 Mad. genera 1 IV Ar Ar

_. xë"gpbG& ie"~iros~~. 11 S genera 1 IV Ar Ar Xenopimstris damii 11 W genera O 1 Ar Xenopimstris polleni 11 1 E genera 1 I Ar Fatculea palliata Ill 11Mad. genera 2 IGV 'Ar Leptopterus viridis I 11,iI I I II Mad. genera 2 IV Ar Ar Leptopterus chabert 111111 1 Mad. genera 2 l Veg Ar Ar Cyanolaniusmadagascarinus 1 1 1 1 1 1 subspecies 2 1-Veg Ar Ar Oriolia bemieri 11 1 E genera O I Ar Eufycemsprevostii 11 1 E genera O IV Ar Hypositta corallimstris 11 1 E genera O I Ar Tylas eduardi 1 111 1 Mad. genera 1 I Ar Dicmms famicatus 111111 1 subspecies 3 I Ar Hartlaubius auratus 111111 1 Mad. species 3 Veg Ar Ploceus nelicoutvi 11111 E species 1 I Ar Ploceus sakalava 111 W species 3 lVeg Ar Foudia madagascariensis 11111111 Mad. species 3 lVeg Ar Foudia omissa Il 11 E species 1 lVeg Ar TOTAL ...... 68 63 77 89 95 59 33 99 360 L. mm

Table 111. Similaritles between sites according to Jaccard. indices above diagonal refer to endemic specles and non-endemic. species; indices beiow thediagonal refer to to non-passerine specles and passerine species.

lsites 1 RN1RN1 PN RS RN1 RS Maso RN1 PN RS PN RS RN1RN1 Zomb lhot Toli RS RN1 Bere W!1 5 4 11 5 21 3 16 12132279 23 10 67 .77 .72 .78 67 .72 .76 .46 .41 .23 .34 .32 .34 .35 .31 .23 .25 .23 .21 .29 .23 lRNlAndohahela 11 (humid) 1 .78 ,913 .96 .85 .88 .92 1.00 .93 .85 .78 .93 .82 .82 .85 .93 67 :78 .74 .70 .82 .78 IRNI 5 1 .74.68 65 51 .54 .60 .29.30.20.35.49.69 .31 .23.23.25.29 .23 .25.19 .21 1 Andringilra 1 .73 .74 69 .71.69.76.71.78.70.73 .62 62 .57.62.54.57.58.71.64 PN4 .77 57 .71.86 61 .71 .81 .32.19.39.52 .29 .20.25.18.22.27.31.33 Ranomafana .89.79 1.00 .88 .92 .92 -96 .85.96 67 .89 .85 .85 .88.96.69.81.77.73.79.81 RS 21 .7i62 .92 69 .63.78.81 .32.18.38.48 .29 .23.22.28.31 .20 .20.26.17 I .87.73.89 .88 .92.96.88.74.89.85 .85 .88 .96.69.81.77.73.79.81 1 1 .78.59.72.67.70.75.78.48 .45 .24 .38 .31 .38.36.30.25 .26 .23.30.22 lgti&ena .84.88.82 .80 .67 .82 .85.85 .91.72.83.64.85 .84 .88 .88 .85 .69 68 .76.81 RS 16 .65.51 .60 .63.64 .70 .71 .36.39 .21.28.26 .31 .33.27.23.23 .19 .16 .23.19 Arnbafovaky .82 58 .80 .78.85 .88.88 38 .88 67 .81 .85 .85.81 .88 .62 .80 .69 65 .78.73 Maso .74.61.81.84.74 .66 .84.46.43.24 .37 .33 .37.38.34.26 .28 .26.21 .31 .26 Masoala .82.76.64 .8i .73.77.65.92.85.81.85.70.84.92.86 69 .77.75 RN1 12 .79 .86 .81 .84 .74 67 .95 .45.40 .23 .34 .31 .35.35.32.24 .26 .24 .20.30.24 Maroieiv .87 .89.77 .67 .86 .86.70.82.70 .74.78.67.93.85 .82 .82 .93 .78 .85 .93 .78 PN 1 .63 .75.49 69 .43.58 .60 .61.46.26.51.37.40.34.32.23.26.24.31.23 Monl.Ambre 61 68 .60 .58.63.64 60 .58.85.78.93.89.921.00.73.85.81.77.82.65 RS 3 .57 .42 .55 .55 .60 .51 .56.34.53.56 .55 .51 A6 .38.44.33.35.41.42 Manongafivo .56.56.59.51.52.54.53 .51 .72 69 .85 .81 .81 .85.84 64 .76 .72 68 .74 .76 PN 2 .50 .36 .41 .50.45.40.47.50.47.42.38.41.39 .41 .48.45.41.42.39.46 lsalo .33.39.32.37.35.32.48.54.78 68 .68.70.78 .71 .63 65 .69.68.75 RS 2 .57.43.58.57.42.63 61 .69 62 .62.72.59 .66 .56.43.46 .39 .43.53.42 Ankarana .73.93.92.89.57,77.70.42.46.48.42.43.47.44 35 .85.89.70.81 RN1 7 .52 .33 .61.46.47.54.51.65.79.71.56.54.57.49.48.41.51.48 .51 Ankarafantsika .43.49.44 .45 .47.47.43.59.75.56.801.00 .88.89 69 .88.77 67 .92.81 RN1 9 .53 .35' .52 .49.56.45.52 .53 .54.47.57 .66 .82.79 65 .51 .54- .47.46 61 .51 Berneraha .44.49.51.44.45.50.47 68 .88 61 .89.88.84.87 59 .77.88 67 .92 .81 Kiri .83.82.75.57.51.54.53.52.45.52.49.38 .53 .74 .58.51 .51 .43 64 .55 Kirindy .48.44.47.46.49.50.46.58.81.50.80.85.84.92.72.88.69.81.92 Zomb .40.43.47.35.51 .50 .51 .56 .49 .64.70 .54.76.71 66 .49.59 64 .64 Zombitse 50 .48.52.52.51.48.49.46 .6i .85.82.77.81.85.73.88.81.84.54.79 lhot .54.53.47.42.41.40.35.38.39.33.40 .60 .57 61 .69.63.64.81 59 lhotry .33 .57.56.59.58.52.55.42.31.34.36.31.32.36 61 .71 .82.77 69 .78 Toli .58.47.48.44.43.39.42.40.39.28.43 .58 .69 69 .80.65.76.77.78 .81 Toliara .38 .44.63.48.37.40.42.37.38.40 67 .66 .71.64 69 .76 .91.88.68.87 RS 23 .48 .35 .59.65.49.56.48.47.36.43.47 66 .71 .71 .81.64.78.79.93 Beza-Mahafaly .31.31.33.29.29.30.30 .29 .83.78.55.57.50.50.52.46.50.35 .71 .95.77 RN1 10 .38.30.40.35.45 .40 .49 .56.59.53.62.45 63 .79.73.61 Tsfmanampekoka .29.30.29 .28 .32 .30 .30 .50.48.56.50.48.39.28 53 .78.73.78.67 .75 Eere .34.52 .45 .38.48.45 .48 .50.53.52 .60 68 .74.77.59.85.76.81.80 .63 .84 Berenty .38 .40 .38.37.42.40.37.48 63 .67.50 .66 .64 .66 59 .83 .81.80.83.92 RN1 ll'(dry) .48.32.43.43.43.36 .47 .48 .51.53.65 64 .71.71.76.87 .69 .84.93.73.79 Andohahela .32.33.31 .31 .35.33.33.31.39 .53 .50 .56 .55 54 .61.59.78.88.95 .83 -88 BlRD COMMUNITIES IN MADAGASCAR 361

Table IV. Distribution of forest species among ecological nichesin different Phytogeographic Regions and Domains (abbreviation as in TableII).

Ecological niches: Domains and RegionsEcologicalDomainsniches: and Feeding Madagascar e hmsa Food habit w sw c 1 Ar ArIP 1614 13 1330 26 9 .30 34 I T Ar 223 232 3 4 1 T T 11 12 2 3 I Ar T 323 341 4 4 Subtotal Insectivores 18 19 23 32 39 16 9 39 45

1, Veg Ar Ar 889 997 79 11 1, Veg T Ar 435 433 15 8 1, Veg T T 111 22 12 2 Subtotal Omnivores 13 12 15 15 14 10 9 16 21

Veg Ar Ar 667 778 38 9 Veg T Ar 111 111 1 1 Veg T T 222 111 1 2 Subtotal Vegetanans 9 9 10 9 9 IO 3 IO 12

nVAr Ar 15 1611 1116 16 4 17 19 nV T Ar 111 111 11 1 nV T T 111 442 14 4 nV Ar T 111 11 11 1 Subtotal small predators 1914 18 22 22 14 7 23 25

Large predators 434 55325 5

Aerial insectivores 666 666 36 6

TOTAL 68 63 77 89 95 59 33 99 114 L. WILME

Table V. Tolerance to forest degradation of Malagasy forestbird species (numbers are numberof species within taxa)

Number of species in forest types: Taxonomic level of endemismundisturbedsecondaryanthropogenicslightly disturbed growth grassland growth disturbed

Species belongingto an endemic family 8 4 O Species belongingto an endemic subfamily 13 11 2 Species belongingto an endemic22 genera 34 6 Sub total...... 55 37 8 1

Other endemic species 30 26 19 14 Non endemic species with endemic23 subspecies 23 21 19 Other non endemic species 6 6 6 6 Sub total...... 59 55 46 39

TOTAL ...... 11454 92 40