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An avifaunal survey of the Trans-Mara Forest, Kenya PDF

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AnavifaunalsurveyoftheTrans-MaraForest,Kenya 61 An avifaunal survey oftheTrans-Mara Forest, Kenya L.A. Bennun The Mau Forest complex forms the largest continuous tract of indigenous forest in Kenya. Collectively its five administrative blocks (the Eastern, Western and South- westernMau,Trans-Maraand01PusimoruForests: seeFig. 1)haveagazettedareaof some 244000 ha, about 14.5 per cent of the total gazetted forest area in the country (Wilson 1988a).However,thefloraandfaunaofthisimportantgroupofforestsremains little-known (Wilson 1988b). While the avifauna of the high (2700-3000 m) and floristically distinct Eastern Mau was described by Sessions (1966, Britton 1979), avifaunalstudieselsewhereappeartobelimitedtoshortlists(onemorethanfortyyears old) of birds from two sites in the South-Western Mau (Toschi 1946, Gichuki et al. 1988). Observercoverageforsquare61 oftheBirdAtlasofKenya,inwhichmostofthe MauForestcomplexlies,isestimatedasbetterthan75percentforonlyoneofthefour quartersquaredegrees (Lewis &Pomeroy 1989). This paper reports the results of bird survey work conducted in the Trans-Mara ForestduringJanuary-February 1990,when somecomparativedatawerealsocollected in the South-Western Mau. The survey forms part of a larger environmental study carried outintheseforests. TheTrans-MaraForest Thisforest,inNarokDistrict,south-westKenya,coversabout35000ha(14percentof the total complex area) in the south-west of the Mau complex. Altitude ranges from about2100min the westto 2400mneartheeastern boundary. Following this shallow gradient, typical moist lowermontane forest grades upwards through a transition zone of 'poor' forest (Kerfoot 1964) into bamboo/forest mixture orpure stands ofbamboo Arundinariaalpinaaboveabout2300m.Withinthegazettedforestboundarypartofthe bamboo zone has been cleared recently to give way to tea plantations, while the characteroftheforestatloweraltitudeshasbeensubstantiallychangedbyextensiveand destructive logging that took place from 1979 to 1986 (J. Howell, pers. comm.). Dominant large timber trees once included species such as Polyscias kikuyuensis, Albizia gummifera and Olea hochstetteri, but few specimens of these or other timber speciesnowremain.TimberremovalhastakenplaceovermostoftheareaoftheTrans- Maraandpartofthe South-Western Mau, upto about 12kmeitherside oftheexisting Olenguruone-Silibwet road. The forest is most degraded near the western boundary, with little high canopy cover. Further east, more tall trees remain and there has been substantial regrowth of non-timber species such as Neoboutonia macrocalyx and Tabernaemontanastapfiana.Removalofthecanopytreesinmanyplaceshasresultedin a dense and tangled undergrowth layer, dominated by Mimulopsis sp. and Piper capense,givingwaytoAcanthuseminensathigheraltitudes.Indampersites,especially valleybottoms,Brillantiasiaiscommonin theunderstory andextensive stands oftree- ferns Cyathea manniana alsooccur. The less-disturbed South-Western Mau appears to have a somewhat different forest structure, although with similar dominant species. In the areas visited, the forest was morelayeredwithadistinctunderstoryofBrillantiasiaandNeoboutonia, andwas gen- erally much more open at ground level due to the better shade. Although rarely com- 62 AnavifaunalsurveyoftheTrans-MaraForest,Kenya 4|Bomel Fig. 1. SketchmapoftheMauForestcomplexshowing net-sitesworkedintheTrans- MaraandSouth-WesternMau plete,thehighcanopywaslessbrokenthanintheTrans-Mara(cf. Kerfoot 1964). Excluding the bamboo zone, forest structure in the Trans-Mara appears rather uni- form, despite gradients in altitude and disturbance. Differentiation between ridgetop, hillslopeand valley foresttypes isprobablypoordueto therelatively smooth topogra- phy,with shallow valleys andbroadridges, andhasbeenfurtherblurredbylogging, Methods SurveyworkwascarriedoutintheTrans-MaraForestReserveover22daysbetween28 January and 23 February 1990. Briefvisits were also made to the South-Western Mau NatureReserve(20-22February)andForestReserve(24-26February)(Fig. 1). Mist-netswereusedateightsiteswithintheTrans-Mara(AtoHinFig. 1),coveringa rangeofaltitudeandforestdisturbance.Aswellasmontaneforest,thesitessampledtwo distinct habitats, tree-fern forest (site G) and bamboo (site H). For ease ofaccess, all sites were close to the alignment of the existing Olenguruone-Silibwet road (Fig. 1). TwoadditionalsiteswereoperatedinthecomparativelyundisturbedforestoftheSouth- western Mau, one at low altitude in the Forest Reserve (Kipteget River, 2020 m) and oneathigh altitudeintheNatureReserve(4kmNEofBostaHill, 2320m; Fig. 1, sites J andIrespectively). Most net sites made use of existing narrow trails in the forest; two used transects recentlycutforvegetationmeasures.Ateachsiteeighttotensmall-mesh,3-mhighnets were set, representing 96-141 m in total length. Wherever possible nets were set in a line with only small gaps between them. Nets were opened for two morning (dawn to about 10:30) and twoevening (16:30 until dusk) sessions atthe first five sites, and for twomorningsessionsonlythereafterbecauseofdeterioratingweather.Attwositesnets AnavifaunalsurveyoftheTrans-MaraForest,Kenya 63 wereoperatedforonlyonefull sessionbecauseofrain.All birdscaughtwereringed. JanuaryandFebruaryareusuallythedriestmonthsintheMauForest(Kerfoot1964), although some rain is always recorded. Conditions in the Trans-Mara were dry on arrivalinlateJanuary,butdeterioratedsteadilyoverthenextmonth.Bythelastweekin February heavy rain occurred every afternoon, often continuing throughout the night andfollowingmorning. Vegetation measures were made at each net site over five quadrats, positioned m alternately5 toleftandrightofthenettrailfromtheendofeverysecondnet(starting with net 1). The percentage cover ofherb (<0.5 m in height), leaflitter and soil were m recordedina 1 x 1 square,andthepercentagecoverofvegetationinfourlayers(0.5- 1 m, 1-3 m, 3-8 mand>8 m) in a3 x 3 m square. Results wereaveragedforeach net site. Vegetation at two sites was not assessed because of bad weather. Disturbance at each sitewasratedsubjectivelyonascaleof (none)to 10(highlydisturbed). Casual observations were made at any time throughout the forest. In addition, to obtain measures ofrelative abundance for canopy species (which are not sampled by mistnets)aseriesoftimedspeciescountswascarriedout(Pomeroy&Tengecho 1986). CountsfollowedthemethodoutlinedbyPomeroy&Tengecho(1986),withtwomodifi- cations: followingpreliminary trials which showedthatfew species wereaddedduring thelastthird ofacount,theduration ofeachcountwasreducedto40min from60min, andonlybirds seen orheardcallingwithin25 moftheobserverwerelisted.Thislatter modification was designed to prevent inflated measures of relative abundance being calculated for conspicuous species (see Pomeroy 1989). For each count, each species was scored according to when it was recorded (4 ifin the first ten minutes, 3 ifin the next ten minutes and so on) and scores were averaged overallcounts to give a simple relative abundanceindex.Thesecountsignored birds movingbelow 5 minthevegeta- tion, and no indices were calculated for inconspicuous mid-level species such as greenbulsthatwereoccasionally seen highinthecanopy. Ten counts were made between 30 January and 16 February 1990 at various points along theforesttrack and various times ofday.- In allcounts I walked slowly along the roadwhilescanningthetrees. Deterioratingweatherforcedtheseriestobecutshort,so the results should be considered provisional: ideally, 15 counts or more should be conducted over a longer period to provide a good picture of a regional avifauna (Pomeroy&Tengecho 1986). Results Speciesrecorded Ninety-two species were recorded within the Trans-Mara forest block (Table 1) of which82(89percent)canbeconsideredforestorforest-edgebirds(Britton 1980).The observationsprovide55 newrecordsfortheBirdAtlasofKenya, ofwhich22arerange extensions. Only one additional species (Broad-billed Roller Eurystomus glaucurus) wasrecordedduringobservation andmist-nettinginthe South-WesternMau. Undergrowthspecies In all, 328 birdswerecapturedin theTrans-Maraand another64in theSouth-Western Mau in 10 000 mh'1 of netting. Capture rates varied between 3.1 to 5.1 birds per 100mIr1 in the forest butwere much lower (1.4/100 mh"1) in the bamboo. Excluding thebamboosite,forwhichno vegetation measures wereavailable,capturerates across sitesweresignificantlypositivelycorrelatedwithvegetationcoverbetween0.5 and 1 m 64 AnavifaunalsurveyoftheTrans-MaraForest,Kenya Table I.Annotatedlistofspeciesrecorded SpeciesarelistedbytheirnumberintheBirdAtlasofKenya(Lewis&Pomeroy 1989). Newrecordsforatlassquares61Cand61Dareindicatedundertheappropriatecolumns asfollows: **,firstrecord; *,firstrecent(post-1970)record;PB,firstprobablebreeding record; B, firstdefinitebreedingrecord. Othercolumns: N^,numbercaughtinTrans- Maraforest;Nsw,numbercaughtinSWMauforest;BZ,bamboozone:+,recorded,-not recorded. 2IZ__ 6_1C _61D_NTO_NS_W BZ Scopidae(Hamerkop) 44HamerkopScopusumbretta Alongroadway. Accipitridae (birdsofprey) 116MountainBuzzardButeotachardus * - - + Especiallycommoninbamboozone. 121 Long-crestedEagleLophaetusoccipitalis - - 122CrownedEagleStephanoaetuscoronatus PP - 135 BlackKiteMilvusmigrans * Phasianidae(quails,francolins) 161 ScalyFrancolinFrancolinussquamatus B - Numididae(guineafowls) 180HelmetedGuineafowlNumidameleagris Occasionalrecordsthroughoutforest. Scolopacidae(sandpipers,etc.) 262Green SandpiperTringaochropus - Pondsalongroadway. Columbidae(pigeons,doves) 325 OlivePigeonColumbaarquatrix - + 326Bronze-napedPigeonColumbadelegorg'uei - 330Red-eyedDoveStreptopeliasemitorquata + Onlyinbamboozone. 334LemonDoveAplopelialarvata ** 9 2 335TambourineDoveTurturtympanistria B ** 9 1 340GreenPigeonTreronaustralis Musophagidae (turacos) 358 Hartlaub'sTuracoTauracohartlaubi * - - + Cuculidae(cuckoos) 365 Red-chestedCuckooCuculussolitarius + 366BlackCuckooCuculusclamosus - 372EmeraldCuckooChrysococcyxcupreus ** ** - - + AnavifaunalsurveyoftheTrans-MamForest,Kenya 65 61C 61D N N BZ ra SW Strigidae (owls) 389AfricanWoodOwlCiccabawoodfordii 392Red-chestedOwletGlaucidiumtephronotum ** Callingshortlybeforedawnonmostmornings. Caprimulgidae (nightjars) 398 MontaneNightjarCaprimulguspoliocephalus Alongroadway. Notcalling. Meropidae (bee-eaters) 444Cinnamon-chestedBee-eaterMeropsoreobates Commonalongroadway;nest-holesinbanks. Phoeniculidae (woodhoopoes) 461 White-headedWood-hoopoePhoeniculusbollei **B Bucerotidae (hornbills) 464BlackandWhiteCasquedHornbill Bycanistessubcylindricus 474CrownedHornbill Tockusalboterminatus Onerecordonlyinhighlydisturbedforest. Capitonidae (barbets,tinkerbirds) 489 Grey-throatedBarbetGymnobuccobonapartei 495Yellow-rumpedTinkerbirdPogoniulusbilineatus 496Yellow-billedBarbetTrachylaemuspurpuratus ** Indicatoridae (honeyguides) 500Black-throatedHoneyguideIndicatorindicator ** Onerecord,calling. Picidae (woodpeckers) 515 Fine-bandedWoodpeckerCampetheratullbergi 519CardinalWoodpeckerDendropicosfuscescens Hirundinidae (swallows) 557 MosqueSwallowHirundosenegalensis Onerecord,highlydisturbedforest. 564BlackRough-wingPsalidoprocnepristoptera PB Oriolidae (orioles) 570MontaneOrioleOrioluspercivali ** Paridae (tits) 583White-belliedTitParusalbiventris 66 AnavifaunalsurveyoftheTrans-MaraForest,Kenya 61C 61D N N BZ TO SW Timaliidae(babblers) **"- 589Pale-breastedIlladopsisTrichastomarufipennis 1 Onecaptured(seeAppendix). 591 MountainIlladopsisTrichastomapyrrhopterum *PB - 19 2 594AfricanHillBabblerAlcippeabyssinica PB - 12 - Campephagidae (cuckooshrikes) 604BlackCuckoo ShrikeCampephagaflava - Fewrecords. 607 GreyCuckoo ShrikeCoracinacaesia * Pycnonotidae (bulbuls) 609CommonBulbul Pycnonotusbarbatus 1 - + 613 Slender-billedGre&nbulAndropadusgracilirostris ** 615 MountainGreenbulAndropadustephrolaemus **pg * 14 5 - 618Yellow-whiskeredGreenbulAndropadus latirostris PB * 29 5 - 626PlacidGreenbulPhyllastrephusplacidus **pg * 29 3 + Turdidae(thrushes) 662White-starredForestRobinPogonocichlastellata B 31 4 + 663 Brown-chestedAletheAlethepoliocephala ** 3 5 - 665 Snowy-headedRobinChatCossyphaniveicapilla **PB - 2 2 - 670RobinChatCossyphacajfra PB - 1 - - 672EquatorialAkalatSheppardiaaequatorialis *PB - 7 8 - 678 Northern OliveThrush Turdusabyssinicus PB - 9 2 - 680AbyssinianGroundThrush Turduspiaggiae **PB - 18 4 - Sylviidae (warblers) 684CinnamonBrackenWarbler Bradypteruscinnamomeus ** * 8 + 685 EvergreenForestWarblerBradypterusbarratti ** 4 . - 703 MountainYellowWarblerChloropetasimilis ** 2 . - 708 BlackcapSylviaatricapilla - 29 2 - 712BrownWoodlandWarblerPhylloscopusumbrovirens PPBB - 4 - - 714ChiffchaffPhylloscopuscollybita ** 715WillowWarblerPhylloscopustrochilus Onerecord. 746BandedPriniaPriniabairdii *B - 16 751 Black-throatedApalisApalisjacksoni - Onerecord. 754GreyApalisApaliscinerea ** 756Chestnut-throatedApalisApalisporphyrolaema 757 Black-collaredApalisApalismelanocephala - - 1 Onerecordinhighlydisturbedforest. 759Black-facedRufousWarbler Bathmocercuscerviniventris PB - 1 Onerecordindensetree-fernforest. Anavifauna!surveyoftheTrans-MaraForest,Kenya 67 61C 61D N N BZ TO SW 769White-browedCrombecSylvietta leucophrys 3 Muscicapidae (flycatchers) 782DuskyFlycatcherMuscicapaadusta - 790White-eyedSlatyFlycatcherMelaenornischocolatina - - 1 799 Chin-spotBatisBatismolitor 803 Black-throatedWattle-eyePlatysteirapeltata PB - 4 809White-tailedCrestedFlycatcher Trochocercusalbonotatus PB - 8 812ParadiseFlycatcherTerpsiphoneviridis - Motacillidae (wagtails,pipits) 829CapeWagtailMotacillacapensis - Alongroadway. 830YellowWagtailMotacillaflava Alongroadway. Malaconotidae (bushshrikes) 847Luhder'sBush ShrikeLaniariusluehderi PB 2 849TropicalBoubouLaniariusferrugineus - - 853 Many-colouredBushShrikeMalaconotusmulticolor ** - - Sturnidae(starlings) 875 Stuhlmann's StarlingPoeopterastuhlmanni 887Waller'sChestnut-wingedStarling ... Onychognathuswalleri - 897 Sharpe'sStarlingCinnyricinclussharpii Nectariniidae (sunbirds) 902Collared SunbirdAnthreptescollaris - 921 NorthernDouble-collaredSunbiidNectariniapreussi - 929 Olive SunbirdNectariniaolivacea - - 13 934Green-headed SunbirdNectariniaverticalis - Zosteropidae (white-eyes) 937YellowWhite-eyeZosteropssenegalensis PB * 18 Ploceidae(weavers) 960BaglafechtWeaverPloceusbaglafecht - Alongroadwayandindisturbedforest. 964Black-billedWeaverPloceusmelanogaster PB 6 965 Brown-cappedWeaverPloceusinsignis PB - Estrildidae (waxbills) 1005 Grey-headedNegrofinchNigritacanicapilla - Onerecord. 68 AnavifaunalsurveyoftheTrans-MaraForest,Kenya 61C 61D N N BZ TO SW 1016AbyssinianCrimson-wingCryptospizasalvadorii - - 13 1 1036Black-crownedWaxbillEstrildanonnula - Alongroadway. Fringillidae(buntings, etc.) ..... 1063Thick-billedSeed-eaterSerinusburtoni - Fewrecords. 1064StreakySeed-eaterSerinusstriolatus Disturbedforestonly,fewrecords. (Spearman's r = 0.71, n = 8, P< 0.05) but not with any other vegetation feature. The only vegetation feature strongly associated with disturbance was canopy cover above 8 m (r = -0.80, n = 8, P< 0.02). A simple measure of species diversity calculated for each site, the Shannon index (Krebs 1978), was also negatively correlated with cover above 8m (r= -0.82, n = 8,P=0.01) but showed no significantrelationship with dis- turbanceperse (r=-0.53,n=9,P>0.15). Itwas difficultto relate the distribution ofundergrowth species tovegetation meas- ures in finer detail. Most species were widely distributed across sites, although some common species such as Brown-chestedAlethe and Evergreen Forest Warbler had an inexplicably patchy distribution. Three species characteristic of disturbed forest and scrub, the Common Bulbul, Robin ChatandWhite-eyed SlatyFlycatcher, werecaught only at the most disturbed and open site (site A). Yellow-whiskered, Mountain and Placid Greenbuls, usually among the most frequently caught species, were almost absent at this site (only oneYellow-whiskered Greenbul captured, none ofany species seen) suggesting they are mainly restricted to the better-preserved forest. The Black- faced RufousWarblerwasclearlyconfinedtotheleastdisturbed areas. Itwascaptured and seen atboth sites in the South-Western Mau (I andJ), where it appeared common and conspicuous in dense shaded undergrowth, but none was seen in the Trans-Mara and onlyonewascaught,inapatch ofdenseundisturbedtree-fern atsiteG. Several species appeared to show altitudinal limitation. Britton (1980) records the Snowy-headedRobinChatandEquatorialAkalatupto2000mand2200mrespectively. IntheTrans-Maratheywerecaughtonlyatthetwolowestsites(Aand B,2200mand 2240m).TheywererecordedagaininlessdisturbedforestintheSouth-WesternMauat Kipteget River (J, 2000 m) but not at Bosta (I, 2320 m). The White-tailed Crested Flycatcherwascaughtateveryforest siteexceptthetwo lowestones(JandA, 2000m and 2200m); however, itis known in KakamegaForestfrom as low as 1650m (Mann 1980). Noneofthese speciesislikelytohavebeenoverlooked atsiteswheretheywere present. Canopyspecies The relative abundance of canopy species, calculated from timed species counts, is shown in Table 2. The three commonest species (Black Rough-wing Swallow, Cinna- mon-chested Bee-eater and Common Bulbul) all forage mainly over or along theroad (wheretheswallowsandbee-eatersalsobreed),sotheseindicesdonotgiveanaccurate pictureoftheirrelativenumbersovertheforestasawhole.Nosuchbiasshouldexistfor Anavifaunalsurvey ofthe Trans-MaraForest,Kenya 69 Table2. The relativeabundance ofcanopyspecies inthe Trans-Maraforestas indicatedby timedspeciescountsalongtheroadway (see textfordetails). The maximumabundance indexis4.0 Abundance index Species 3.3 BlackRough-wing Swallow 2.8 CommonBulbul 2.6 Cinnamon-chestedBee-eater 2.5 Chestnut-throatedApalis 2.0 Yellow-rumpedTinker-bird 2.0 Blackcap 1.5 YellowWhite-eye 1.4 OliveThrush 1.4 Chin-spotBatis 1.2 MontaneOriole 1.1 White-headedWoodHoopoe 1.0 DuskyFlycatcher 0.9 Northern Double-collared Sunbird 0.8 Sharpe's Starling 0.8 GreyCuckoo Shrike 0.7 Red-chestedCuckoo 0.7 White-eyed SlatyFlycatcher 0.6 Fine-bandedWoodpecker 0.6 GreyApalis 0.5 BrownWoodlandWarbler 0.4 Yellow-billedBarbet 0.4 Slender-.billed Greenbul 0.3 Collared Sunbird 0.3 Brown-cappedWeaver 0.2 Hartlaub'sTuraco 0.2 Waller's Chestnut-wingedStarling 0.2 Emerald Cuckoo 0.2 White-browedCrombec 0.2 Chiffchaff 0.2 Green-headedSunbird 0.1 Grey-throatedBarbet 0.1 Black-frontedBushShrike the remaining species. The results indicate the advantage of even a short sequence of counts over simple observation: for instance, Red-chested Cuckoos were heard calling almost constantly, but their true numbers (as the counts suggest) must be relatively small.Notethatthreespecies,YellowWhite-eye,Blackcap,andNorthernOliveThrush, werecommonlyrecorded bothincounts andin mist-nets (Tables 1 and2). 70 AnavifaunalsurveyoftheTrans-MaraForest,Kenya Discussion In spite ofits disturbed state, the Trans-Mara forest still supports a diverse avifauna. Elevenofthe82forestspeciesrecordedthere(15percent)aredistinctlywesternintheir affinities, namely the Red-chested Owlet, Grey-throated Barbet,Yellow-billed Barbet, Mountain Illadopsis, Snowy-headed Robin Chat, Equatorial Akalat, Banded Prinia, Black-faced Rufous Warbler, Luhder's Bush Shrike, Stuhlmann's Starling and Black- crownedWaxbill.TheRed-chestedOwlethasbeenrecordedintheKikuyuEscarpment & forest (Taylor Taylor 1988) and the Mountain Illadopsis in the Aberdares (Britton 1980),butotherwisenoneofthesespeciesisknowneastoftheRiftValleyinKenya.In terms ofglobalconservation, all these species have populations in otherparts oftheir range,chieflyinUgandaandeasternZaire.However,recentinformationontheirstatus is lacking and several areprobably uncommon throughout theirrange (D.E. Pomeroy, pers. comm.). In addition, these western species are of special interest within Kenya because the other forests in which they occur are small, fragmented and increasingly degraded. Theapparentp esenceofPale-breastedIlladopsisintheTrans-Maraissurprisingand needs confirmation (see Appendix). This species has a disjunct distribution, with the nominate race reaching Kenya at Kakamega and South Nandi while the race distans occupiesforests700kmawayontheeasternarcmountainsinTanzania(Britton 1980). Significantly,thespecieshasrecentlybeendiscoveredontheforestislandof01Doinyo Orok, almost half-way between these two populations (Bennun et al. 1986). The 01 Doinyo Orok birds, like the individual caught in the Trans-Mara, appeared closer to distans than to the nominate race. The 01 Doinyo Orok record is at 1800 m and the altitudinal limit elsewhere in Kenya is at about 1700 m (Britton 1980, Mann 1980), m comparedto2240 fortheTrans-Mararecord.However,thereisasinglerecordinthe Cherenganis at 2300 m (Britton 1980). This, together with the Trans-Mara record, suggests both that this species may be frequently overlooked and that it may range considerablyhigherthan hasbeenthought. Some 23 additional forest or forest-edge species previously recorded from atlas square61C(Lewis&Pomeroy 1989), whichincludesmuchoftheTrans-Maraandthe South-WesternMau,werenotfoundduringthepresentsurvey.Additionalworkwillno doubt addmany ofthese totheTrans-Maralist, particularly thosethatareinconspicu- ous(e.g.MoustachedGreenTinker-bird,Purple-throatedCuckoo-shrike)orthatvaryin abundance through seasonal altitudinal migration (e.g. Narina'sTrogon). Others, espe- cially those whose mainranges are athigherorloweraltitudes, orthatareknown only fromoldrecords,mayneverhaveoccurredintheTrans-Maraitself.Therearenonethe- lesssomesurprisingabsences,includingCrestedGuineafowl(itsplaceapparentlytaken by the Helmeted, usually a woodland bird), African Goshawk and Doherty's Bush Shrike.ThepresenceofRed-frontedParrotwould alsobeexpected, althoughithasnot yet been recorded from this atlas square. It is clear that habitat modification has had some effect on species composition, with the encroachment ofedge species along the road and western margins; such infiltrationprobably accounts forthenegativecorrela- tion between undergrowth species diversity and highcanopycover. Mostundergrowth species appearto have adapted surprisingly well to the altered forest structure, butthe effects on canopy birds are less clear-cut; the possibility of local extinction in some cases due to forest destruction cannot be ruled out. Further study of the entire Mau complex should illuminate these issues, which are ofconsiderable importanceforbird conservationinthisandotherEastAfricanforests

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