La forêt de Niaouli se trouve sur le territoire d'une station de recherche agricole dans le Sud du Bénin. Protégée depuis 1997, la forêt semi-décidue de 150 ha (110 ha de forêt de plateau et 40 ha de forêt de bas-fonds) forme un îlot forestier dans la région. Des recherche ornithologique ont été effectuée depuis 1997 pour établir un inventaire et d'étudier la migration intra-régionale. Au total, 166 espèces d'oiseaux ont été identifiées, dont 14 nouvelles espèces le Bénin: l'Engoulevent à épaulettes noires Caprimulgus nigriscapularis, le Martin-pêcheur à ventre blanc Corythornis leucogaster, le Barbican chauve Gymnobucco calvus, le Barbion grivelé Pogoniulus scolopaceus, l'Indicateur pygmée Prodotiscus insignis, l'Indicateur tacheté Indicator maculatus, le Bulbul à queue blanche Baeopogon indicator, la Camaroptère à sourcils Camaroptera superciliaris, l'Erémomèle à tête brune Eremomela badiceps, l'Hyliote à dos violet Hyliota violacea, l'Akalat à ailes rousses Illadopsis rufescens, la Mésangette rayée Pholidornis rushiae, le Gonolek fuligineux Laniarius leucorhynchus et Pyréneste ponceau Pyrenestes ostrinus.
Benin lies in the Dahomey Gap, a region characterised by relatively low annual rainfall (800-1,400 mm) and that savanna reaches almost to the coast, with an almost complete lack of the tropical forest that is a feature of coastal zones in adjacent countries. Until the 1970s, how-ever, the region between the coast and up to 150 km inland still held relatively large patches of tall forest, typical of the Guinea-Congo vegetation zone . High human population in the tropical south, where densities reached 250-416 per sq km , has resulted in virtually all of this forest being cleared for agriculture [5, 27]. Only small 'islands' remain: typically sacred areas, small privately owned holdings, and some state-owned areas consisting of plantations and small parcels of secondary forest.
In 1997, the author initiated a forest protection programme in collaboration with the Dutch Embassy, CBDD (Centre Béninois pour le Développement Durable) and NC-IUCN (the Netherlands Committee of the International Union for Conservation of Nature). This led to the protection of one of the remaining forest islands, at Niaouli. The main objective is the conservation of extant natural forest, as well as research on flora and fauna for environmental education and ecotourism.
Niaouli is situated c70 km north of Cotonou (06°44'N 02°09'E). The forest belongs to the national agricultural research station Niaouli, Attogon, and consists of a plateau (dry) forest and a wetter bas-fond (humid) forest covering 150 ha in total. A reforestation belt (15 ha) protects the forest against bush fire spreading from adjacent agriculture, and c20 ha of degraded forest have been reforested with indigenous trees, including flowering species, eg Milicia excelsa and Cassia siamea, to enable bee-keeping activities. Three local rangers enforce a hunting and cutting ban. Other measures have been designed to compensate local villagers who previously used the area as a source of firewood and for hunting. Two ecological trails, one through each forest type, as well as an observation tower on the plateau, were constructed to facilitate education on environmental and natural resources.
Since 1997, several inventories have documented the flora and fauna. A detailed checklist of the avifauna is being prepared through regular field observations and mist-netting, with the aim of not only identifying which species use the forest but also their population densities.
In 2000, an additional small research project was established in five other forests in southern Benin. In each, an inventory is being prepared using the same methods as above, and birds are ringed with the help of the Institut für Vogelforschung 'Vogelwarte Helgoland' (Wilhelmshaven, Germany). One objective is to observe intra-regional migration of resident birds, between the remaining forest islands. For example, a Western Olive Sunbird Cyanomitra obscura ringed at Niaouli on 26 October 2001 was caught, on 13 February 2002, at Pobè (06°58'N 02°40'E), c80 km north-east of Niaouli.
The Benin literature
Such research will also contribute to our still limited knowledge of the Benin avifauna. In comparison with neighbouring countries, relatively little has been published on the ornithol-ogy of Benin, and as recently as 1993 evidence was available for the occurrence of only 425 species in the country , although more recent work, including the present contribution, has increased this total to 570 species (Claffey pers comm).
The early literature is composed of contributions by specimen collectors [13, 24], while several administrators and missionaries made contributions in the colonial period [7, 9, 14]. These were relatively meagre, however, and do not compare with those for Togo, particularly in the early colonial period, when an important contribution was made to the knowledge of that country's avifauna . In the post-colonial period there have been occasional studies and surveys, most notably Green & Sayer's survey of Arli and Pendjari National Parks . Holyoak & Seddon  provided distributional notes from a brief visit to the country, while Claffey's survey of the Bétérou area, in 1987-1995 , covers areas much further to the north, including the classified forests of Ouari Maro, Monts Kouffé and the Ouémé Supérieur (08°30'-09°12'N 02°00-02°16'E), as well as additional notes from the Borgou and other areas. He also published several short notes during the 1990s.
Of forests in the south, Anciaux  inventoried the Allada Plateau, including Niaouli, in 1991-1994, recording 124 species. A preliminary inventory of Lama forest (06°55'-07°00'N 02°04'-02°12'E) was published by Waltert , recording 106 species during 31 field days. More recently, Anciaux has published a study of intra-African migrants from work on the Allada Plateau and in the Lama depression . Several unpublished reports, notably by Miriam Langeveld, who worked in the far north, have also been utilised in preparing the revised list of Benin birds (Claffey pers comm).
For the purposes of the research the forest was considered to comprise two distinct sections. While there were many similarities in the avifauna of the two, some interesting differences were observed. A total 166 species was found in Niaouli on 125 field days (see Appendix 1) during January 1997 to May 2002. Of these, 151 occurred in the plateau forest, while 105 species were observed in the bas-fond forest.
This forest covers c110 ha, including 25 ha of reforestation. Many of the semi-deciduous trees lose their leaves during the dry season (Fig 1). These consist of nine families and 13 species, eg Ceiba pentandra (Bombacaceae), Triplochiton scleroxylon (Sterculiaceae), Antiaris toxi-caria and Milicia excelsa (Moraceae), and Dialium guineense (Caesalpiniaceae). Typical bird species of the upper storey are: African Cuckoo Falcon Aviceda cuculoides, African Harrier Hawk Polyboroides typus, Green Turaco Tauraco persa, African Green Pigeon Treron calva, Blue-throated Roller Eurystomus gularis, Yellow-mantled Weaver Ploceus tricolor, Red-headed Malimbe Malimbus rubricollis and Naked-faced Barbet Gymnobucco calvus. Under-storey inhabitants include Brown Illadopsis Illadopsis fulvescens iboensis, White-throated Greenbul Phyllastrephus albigularis, Grey-headed Bristlebill Bleda canicapilla and Chestnut Wattle-eye Platysteira castanea.
This forest is irrigated by three wells permitting evergreen vegetation throughout the year (Fig 2). It covers an area of c40 ha, including 10 ha of reforestation. The tree vegetation is much more diverse than in the plateau forest, consisting of 28 families and at least 63 species, eg Symphonia globulifera (Clusiaceae), Cleistopholis patens (Annonaceae), Cola gigantea, C. millenii and C. nitida (Sterculiaceae), Musanga cecropioides (Cecropiaceae), Piptadenias-trum africanum (Mimosaceae) and Entandrophragma angolense (Meliaceae) which in Benin occurs only in Niaouli. Differences in the vegetation are reflected in the avifauna. Typical species of the canopy include: Red-thighed Sparrowhawk Accipiter erythropus, Black-headed Oriole Oriolus brachyrhynchus, Chestnut-winged Starling Onychognathus fulgidus and Buff-throated Apalis Apalis rufogularis, while in the understorey are Little Greenbul Andropadus virens, Western Olive Sunbird, Blue-billed Weaver Spermophaga haematina and Blue-billed Malimbe Malimbus nitens.
Discoveries and significant records
Of the 166 species observed (Appendix 1), 14 are additions to the preliminary Benin checklist (Claffey pers comm) and several others concern species for which there was little previous evidence. The first evidence of breeding in the country was noted for a number of species. Status in neighbouring Togo and Nigeria is taken from the respective checklists [10, 16]. Among the most interesting records are the following.
Red-thighed Sparrowhawk Accipiter erythropus
Uncommon resident in Benin (Claffey pers comm). Previously known only from January and April records in Arli and Pendjari Parks . Several records at Niaouli including a female mist-netted on 5 October 2000 (wing length 174 mm, tarsus 43 mm, bill 12.5 mm, weight 133 g; Fig 3). The absence of white uppertail spots indicate the nominate race as would be expected (W S Clark pers comm). Green reported 'a very doubtful record of nesting' at Natitingou in 1978 (Claffey pers comm). I observed material being collected for a nest c40 m above ground, on 6 March 1999, while the male was observed bringing prey to the nest for the female, on 13-15 March 1999.
Black Sparrowhawk Accipiter melanoleucus
Uncommon to rare resident that was first recorded recently . Only one previous record in the Forêt Classée de la Lama in 199425. Recorded five times at Niaouli (in January 1997, August 1998 [three records] and March 2002) and twice at Lokoli marsh forest (07°03'N 02°16'E), in September 2001 and a juvenile on 25 January 2002.
Thick-billed Cuckoo Pachycoccyx audeberti
Rare and patchily distributed in West Africa . The only previous Benin record was one identified by call in the Forêt Classée de l'Ouémé Supérieur (Claffey pers comm). In Niaouli, recorded in June 1998, February and March 1999, November 2001, and March-May 2002. A noisy bird, identified in flight by its slow wing action, and confirmed by its very characteristic vocalisation. Rare resident in Togo and recorded in Nigeria.
Black-shouldered Nightjar Caprimulgus nigriscapularis
Previously unknown in Benin, and considered uncommon to rare in West Africa . Recorded at four different forests in southern Benin in January-March 2002. In Niaouli, the species was identified by voice three times, usually at sunset and dawn. During a full moon it was heard calling throughout the night, at intervals of 8-10 seconds, and once two individuals called simultaneously. One was mist-netted on 24 February 2002, in a humid area of Lokoli (wing length 149 mm, tarsus 14 mm, bill 13.5 mm, tail 124 mm, weight 42 g; Fig 4). Not uncommon in Nigeria and a rare resident in Togo.
White-bellied Kingfisher Corythornis leucogaster
Not previously recorded in Benin, and an uncommon to scarce forest resident elsewhere in West Africa . One record near the plateau on 11 February 1997. Separated from Malachite Kingfisher C. cristata by its white belly and the forest habitat , it was perched on the lowest branch of a teak tree at the limit between forest and adjacent agricultural used land. Rare resident in Togo and an uncommon resident in Nigeria.
Naked-faced Barbet Gymnobucco calvus
Not previously recorded in Benin. Gregarious and very common, with several large colonies in Niaouli. An easily distinguished, dull, thick-billed bird, it often exhibits woodpecker-like behaviour when searching for insects in dead trees. Common to locally abundant resident in Togo and Nigeria.
Speckled Tinkerbird Pogoniulus scolopaceus
Not previously recorded in Benin but probably overlooked as its secretive behaviour makes it difficult to observe in the understorey. Sixteen records in 1997-2001, with singles mist-netted on 21 February 1999, 20 September 2000, 26 October 2001 (wing length 54 mm, tarsus 14 mm, bill 14 mm, tail 31 mm, weight 16 g) and 16 May 2002 (wing length 54 mm, tarsus 13 mm, bill 15.5 mm, tail 30 mm, weight 15 g; Fig 5). Common resident in Togo and Nigeria.
Cassin's Honeybird Prodotiscus insignis
Not previously recorded in Benin. Inconspicuous and secretive, and thus easily overlooked. Nine observations: Aril 1997, June and August 1998, February 1999, November 2000 and March 2001. Identification was based on behaviour, shy and flycatcher-like , small pointed bill and remarkable white outertail feathers. Rare resident in Togo and an uncommon resident in Nigeria.
Spotted Honeyguide Indicator maculatus
Not previously recorded in Benin. One, in June 1998, within scrub forest adjacent to the tall forest of Niaouli. A dark olive-green honeyguide, identification was based on the remarkable stripes on the belly and white undertail feathers with clearly visible dark barring. Rare resi-dent in Nigeria and an uncommon resident in Togo.
Honeyguide Greenbul Baeopogon indicator
Not previously recorded in Benin. Three records (9 January and 20 February 1997, and 9 April 1998) within mixed-species flocks, and a lone observation on 12 August 1998. All were in the lower storey. Similar to a honeyguide but more bulbul-like, the identification was based on the white outertail feathers while the white eye was noted on two individuals. Not uncommon resident in Togo and Nigeria.
Red-tailed Greenbul Criniger calurus
First recorded in Benin from Niaouli on 20 February and 5 March 19991. Another was mist-netted on 28 November 2000. The species is very common in Lokoli marsh forest. Not uncommon resident in Togo and a common resident in Nigeria.
Yellow-browed Camaroptera Camaroptera superciliaris
Not previously recorded in Benin. Six records (14 January, 20 May, 21 June and 30 July 1997, and 3 June and 18 August 1998). Identification of this small, short-tailed species was based on the yellow-olive upperparts, dull white underparts and obvious yellow supercilium. Uncommon resident in Togo and Nigeria.
Rufous-crowned Eremomela Eremomela badiceps
Not previously recorded in Benin. Three records, on 4 and 12 August 1998, and 20 February 1999. Identification was based on habitat, mid-storey of the forest , typical insectivorous behaviour, the white throat, black breast-band and grey-white belly. Not uncommon resident in Togo and Nigeria.
Violet-backed Hyliota Hyliota violacea
Not previously recorded in Benin. A male was seen at close range from the observation tower on 15 May 2002, and identified by its remarkably dark upperparts, tail and head, white belly, and very active behaviour in the mid-storey, where it fed horizontally on the underside of leaves. Uncommon resident in Togo. The record of this subspecies (nehrkorni) is the easternmost to date.
Rufous-winged Illadopsis Illadopsis rufescens
Not previously recorded in Benin. One was mist-netted on 7 June 2001 (wing length 77 mm, tarsus 26 mm, bill 18 mm, tail 60 mm, weight 37 g; Fig 6). Despite the resemblance to Puvel's Illadopsis I. puvelli, which also occurs in Niaouli, the mensural data and photographs permitted its identification as I. rufescens (L D C Fishpool pers comm). Listed as Near Threatened , it is a rare resident in Togo, with only one record. The present record extends its range further east.
Sabine's Puffback Dryoscopus sabini
Previously known in Benin only from Brunel's specimen in MNHN, which has been identified as D. s. sabini . Two records, on 13 March and 20 June 2001. Both were males, the first was within a large mixed-species flock in the mid-storey and the second was searching for insects in the lower canopy. They were distinguished by the obvious white belly and rump, and black head, back and tail. Uncommon resident in Togo and Nigeria.
Sooty Boubou Laniarius leucorhynchus
Not previously recorded in Benin. Two singles on 27 March 1999. This is the only bush-shrike that is all black, with a large black bill and black legs, and that is found in lowland forest . Behaviour is closer to that of a true shrike. Few records in Nigeria and a rare resident in Togo, with only one record.
Tit-Hylia Pholidornis rushiae
Not previously recorded in Benin. Ten records: in February (two), April, May (two), June and July 1997, and June and August (two) 1998. This tiny finch-like bird was identified by its pale brown, finely streaked head, throat and breast, and brown upperparts. Often in small groups of 4-6 individuals. Locally not uncommon resident in Nigeria, and a rare resident in Togo.
Black-bellied Seedcracker Pyrenestes ostrinus
Not previously recorded in Benin. Although Mackworth-Praed & Grant  extrapolated that it occurred in the country, no documentation was available. One was mist-netted, a female on 10 February 2001 (wing length 63 mm, tarsus 19 mm, bill 15.5 mm, tail 49 mm, weight 21.5 g; Fig 7). Uncommon resident in Togo and Nigeria.
How to reach Niaouli
The forest is situated c1 hour north of Cotonou and 5 km from Allada, near Attogon. Follow the sign for the Centre Regional de Recherche Agricole, Niaouli for 3 km. The research station is on the left. Register at the small visitors centre. Reliable guides with good knowledge of the project, as well as the flora and fauna, are available, and it is possible to stay overnight.
F. Toornstra, of the Ambassade Royale des Pays Bas (Netherlands), in Benin, encouraged me to commence the project. Financial support was provided by the Ambassade Royale des Pays Bas in Benin, NC-IUCN and CBDD. J. Kamstra and G. Agbangla are especially thanked for their support. For scientific advice I am indebted to Dr L.D.C. Fishpool (BirdLife International), W. S. Clark (Raptours) and Prof F. Bairlein (Institut für Vogelforschung 'Vogelwarte Helgoland'), and T. Lougbegnon for his assistance during the field work. Patrick M. Claffey provided considerable assistance in editing the second draft, and provided important additional data and references.
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