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M S AMMALIAN PECIES No. 784, pp. 1–7, 3 figs. Pipistrellus nanus. ByCori L. Lausen and Robert M. R. Barclay Published 20 December 2005 by the American Society of Mammalogists Pipistrellus nanus (Peters, 1852) P. nanulus, which hasa sub ‘‘sickle-shaped’’traguswithrounded outermarginanda semicircularantitragus(Rosevear1965). African Banana Bat Although anterior upper premolar is very small, it occurs in allP.nanusandisabsentinmost,butnotall,Eptesicus(Koopman Vespertilio pipistrellusvar.africanusRu¨ppell,1842:156.Typelo- 1975; Rosevear 1965). Lamboidal crests of skull are present, but cality‘‘Shoaprovince,Ethiopia.’’ not prominent, in P. nanus (Fig. 2) and do not form ‘‘backwardly Vespertilio nanus Peters, 1852:63, pl. 16, Fig. 2. Type locality produced helmet’’ present in other Pipistrellus (Rosevear 1965: ‘‘Inhambane[Mozambique],248Lat.’’ 267). VesperugopusillulusPeters,1870:124.Typelocality‘‘Loango,An- gola.’’ GENERAL CHARACTERS. Pipistrellus nanus is one of Vesperugo stampflii Jentink, 1888:54. Type locality ‘‘Farmington thesmallestmammals(Rosevear1965).Earsareroughlytriangular River,Liberia.’’ withouteredgesslightlyconvexandtipspointed(Smithers1983). Vesperugo pusillus Noack, 1889:216, pl. V, Fig. 2. Type locality Antitragal lobe at base of concave margin is a ‘‘pronounced sub- ‘‘Boma,nearmouthoftheCongoRiver,Zaire,’’5Democratic rectangular flap, largely free of the main margin of the ear’’ (Ro- RepublicoftheCongo. sevear 1965:265). Tragus is ‘‘hatchet-shaped’’ with straight inner Vesperugo pagenstecheri Noack, 1889:220, pl. V, Fig. 3. Type lo- margin and outer margin consisting of 2 ‘‘almost straight edges cality‘‘Netonna,Congo,WestAfrica.’’ meeting at an obtuse angle’’ with ‘‘little or no trace of any lobule Pipistrellus minusculus Miller, 1900:647, Fig. 43. Type locality at the base of the outer margin’’ (Rosevear 1965:265). Dorsal fur ‘‘MountCoffee,Liberia.’’ is dark at base with tips yellowish golden-brown or deepreddish- PipistrellusculexThomas,1911:458.Typelocality‘‘Kabwir,North- brown.Ventralfurisdarkatbase,buttipsaremuchlighter,some- ern Nigeria.’’ times grayish or whitish-brown. Adult specimens from Congo are Pipistrellus nanus australisRoberts,1913:67.Typelocality‘‘Port darkbrowndorsallywithfaintreddishcast,anddullandpalebuffy St. Johns, South Africa,’’ not Pipistrellus hesperus australis brownventrally(Allen1917).Colorofadultsvarieswithgeographic Miller,1897. location;specimensfromeastcoastofAfricawerepaleronunder- Pipistrellus abaensis J. A. Allen, 1917:442. Type locality ‘‘Aba, sidethanWestAfricanspecimens(AllenandCoolidge1930),and OrientalProvince,Zaire,’’5DemocraticRepublicoftheCon- specimensfromMalawiweremuchdarkerthanthosefromsouthern go. Tanzania and Kenya (Lawrence and Loveridge 1953). Variationin PipistrellusfourieiThomas,1926:288–289.Typelocality‘‘Ukualu- color also occurs between individual adults and is not influenced kasi,N.W.Ovamboland’’5Namibia. by gender (Lang and Chapin 1917a). Immature specimens are Pipistrellus nanus: Allen, 1939a:95. First use of current name much darker than adults and, in some cases, nearly black (Allen combination. 1917). Wings and interfemoral membranes are brown (Smithers PipistrellusafricanusmeesteriKock,2001:129–130.Replacement 1983).Somespecimensshowtracesofawhiteborderonhindmar- name forPipistrellusnanus australisRoberts,1913:67. ginofwing(Aellen1952; Allenand Coolidge1930). Neoromiciananus:Kearneyetal.,2002:68. Namecombination. Somemalesandfemalespossesstailglands;theseovalstruc- turesare1.2–3.8mmlongandarelocatedonbothsidesoftailon CONTEXT AND CONTENT. Order Chiroptera, suborder anteriorquarterofinterfemoralmembrane(O’Shea1980).Agroup Yangochiroptera (Van Den BusscheandHoofer2004),familyVes- of African banana bats in Kibwezi, Kenya, had these glands pertilionidae, subfamily Vespertilioninae, genus Pipistrellus, sub- (O’Shea1980),whileintheIvoryCoast,someindividualshadthem genusPipistrellus.SpeciesnamenanusPeters,1852isantedated andothersdidnot(HappoldandHappold1996).AlthoughP.nan- byP.africanusRu¨ppell,1842(Koopman1975).Althoughseniority usisconsideredasinglespecies(Koopman1993),2speciesmight of africanus was identified by Koopman (1975), this senior bino- bedifferentiatedbypresenceorabsenceoftheseglands(Happold men isseldom used (Dumontetal. 1999). Asmostliteratureuses and Happold1996). thebinomenP. nanus,Happold(2003)madeaformalapplication Measurements(range,inmm,bothsexes,n$7)forP.nanus (Case3240,InternationalCommissiononZoologicalNomenclature) to conserve use of nanus and suppress use of africanus. A declaration of nonsupport for this suppression was made by Kock (2004). Seven subspecies of Pipistrellus nanus are recognized (Bronneretal.2003; Koopman1994;P. nanus5P. africanus): P. n. africanus (Ru¨ppell, 1842:156), see above (abaensis Allen, 1917 issynonym). P. n. culexThomas,1911:458, seeabove. P. n. fourieiThomas,1926:288–289, seeabove. P.n.heliosHeller,1912:3.Typelocality‘‘MerelleWater,30miles southofMountMarsabit,Kenya.’’ P. n. meesteriKock,2001:129–130,seeabove(australisRoberts, 1913 issynonym). P.n.nanus(Peters,1852:63,pl.16,Fig.2),seeabove(pusillulus Peters1870,pusillusNoack1889,pagenstecheriNoack1889 aresynonyms). P. n. stampflii (Jentink, 1888:54), see above (minusculus Miller, 1900 issynonym). DIAGNOSIS. Pipistrellus nanus(Fig. 1)hasbicoloredhair FIG.1. PhotographofanadultPipistrellusnanusfromSeng- throughout pelage, not just ventrally, as in P. nanulus (Rosevear waWildlifeResearchStation188109S,288139E,Zimbabwe,January 1965). Shapes of tragus and antitragus distinguish P. nanus from 1982. UsedwithpermissionofthephotographerM.B. Fenton. 2 MAMMALIANSPECIES 784—Pipistrellusnanus FIG. 3. Geographic distribution of Pipistrellus nanus. Sub- speciesare1,P.n.africanus(EthiopiatoDemocraticRepublicof theCongo,mostlyinhighlands)asdelineatedbyshadinganddark outline;2,P.n.culex(NigeriaandGhana);3,P.n.fouriei(southern Angola, western Zambia, northern Namibia); 4, P. n. helios (southernSudanandsouthernSomaliatonorthernTanzania);5,P. n. meesteri (Eastern Cape, South Africa); 6, P. n. nanus (Demo- craticRepublicoftheCongoandTanzaniatoKwazulu-Natal,South Africa); 7, P. n. stampflii (Ivory Coast to Sierra Leone). Dashed linedelineatesoverlapof4and6.P. nanusispresentintextured areaswithoutnumbers (x), includingPemba,Zanzibar,andMada- gascar, but subspecies are not defined in these areas. Map used withpermissionofJ.D.Skinner,andadaptedfromitsoriginalform (Skinnerand Smithers1990). mountain region than in the rest of Sudan (Koopman 1975). P. nanushasbeenrecordedwithameanforearmsize,29cm(male) and,31cm(female)inthefollowingareas:southernSudanoutside of the Imatong mountain region (Koopman 1975), Niger (National Park W—Poche´ 1975), Kenya (Kibwezi—O’Shea 1980), Liberia (Miller 1900), Guinea (Lopes and Crawford-Cabral 1990), Nigeria FIG.2. Dorsal,ventral,andlateralviewsofcraniumandlat- (female data only—Bergmans 1977; Thomas 1911), and Senegal eralviewofmandibleofanadultfemalePipistrellusnanus(Royal (femaledataonly—Verschuren1982).P.nanushasbeenrecorded OntarioMuseum71310,8.5kmfromGargenonGaloleRoad,Ken- with amean forearm size$29 cm(male)and$31cm(female)in ya). Condylocanine length is 10.08 mm. Used with permission of thefollowingareas:northeasternDemocraticRepublicoftheCongo thephotographersM.B. Fentonand E. Bernard. (Allen 1917), Equatorial Guinea (Jones 1971), Sudan in the Ima- tongmountainregion(excludingLokwi—Koopman1975),Zimbab- we(SmithersandWilson1979),andMozambique(LopesandCraw- in West Africa are: length of forearm, 25–32.5; length of tail,24– ford-Cabral1990). Extensivemeasurementsareavailablefor1fe- 41; length of tibia, 10–13.5; length of 3rd metacarpal,25.5–31.5; maleand1malefromtheKikwitarea,BandunguProvince,south- length of 5th metacarpal, 25–29.5; greatest length of skull, 11– westernDemocraticRepublicoftheCongo(058139S–188499E—Van 12.5; mastoid breadth, 6.5–7.0; length from canine to 3rd molar, Cakenbergheetal.1999). 3.6–4.3; width between 1st molars, 4.2–5 (Rosevear 1965). Mea- surements (range, in mm or g, sex and sample size notstated)for DISTRIBUTION.AfricanbananabatsoccurthroughoutAf- P.nanusinEastAfricaare:lengthofheadandbody,36–40;length ricasouthoftheSaharaDesert,exceptinaridzonesofNamiband of tail, 24–41; length of forearm, 25–32.5; mass, 2–4 (Kingdon Kalaharideserts(Fig.3;SkinnerandSmithers1990).Recordsare 1974). Females are slightly larger (LaVal and LaVal 1977). Mea- fromAngola,DemocraticRepublicoftheCongo,IvoryCoast,Ken- surements (mean, range in parentheses, in mm or g) for 10 males ya,Malawi,northeastMozambique,Somalia,SierraLeone,southern and 10 females, respectively, from Zimbabwe are total length, 77 Sudan,Tanzania(includingPembaandZanzibar),Uganda,Zambia (72–82),80(74–84);lengthoftail,36(34–38),36(36–38);length (Koopman 1994; Smithers 1983), Cameroon (Aellen 1952), Equa- of ear, 11 (10–11), 11 (11–12); length ofhind foot, 6 (5–7), 6(5– torial Guinea (Jones 1971), Ethiopia and Mali (Hayman and Hill 7);lengthofforearm,32(30–32),32(31–32);mass,4.0(3.0–4.0), 1971),Gabon(Brosset1966),Ghana(Jeffrey1975),Guinea(Lopes 4.0(3.0–4.0—SmithersandWilson1979).Lengthofforearmlength and Crawford-Cabral 1990), Liberia (Allen and Coolidge 1930), increases from west to east from Sierra Leone to southeastCame- Madagascar (Dorst 1947), southern Niger (Poche´ 1975), Nigeria roon(Rosevear1965),andlargerindividualsoccurintheImatong (Bergmans1977;Thomas1911;Vielliard1974),andSenegal(Ver- 784—Pipistrellusnanus MAMMALIANSPECIES 3 schuren1982).P.nanusisrestrictedtonorthernpartsofsouthwest ONTOGENYANDREPRODUCTION.Malereproductive Africa, including northern Namibia and northern Botswana tract consists of paired testes and associated epididymides,which (throughouttheOkavangoDeltaandsouthtoLakeNgami).P.nan- move from a position in lower abdomen during spermatogenesis. usiswidespreadinManaPoolsNationalParkandSengwaWildlife Cauda epididymides extend several millimetersbeyond distalend Area, Zimbabwe (Rautenbach and Fenton 1992), but is not found of testis when full of spermatozoa and are sheathed in a black, in parts of the dry west and south of the Zambezi River in Moz- pigmented membrane, the tunica vaginalis. Accessory glands in- ambique.P. nanushasbeenrecordedinthenorthernandeastern cludepairedseminalvesiclesandacompoundprostate.Ampullary parts of South Africa (Rautenbach and Fenton 1992; Smithers and urethral glands are absent. During earlyspermatogenesis,ep- 1983). Nofossilsareknown. itheliumofseminiferoustubulesis2cellsthick,butprogressively thickensto4cellsbyendofwetseason.Developmentofspherical FORM AND FUNCTION. Thumb pad attaches to flexor spermatidsisfollowedbyproductionofspermatozoabeforestartof side of 1st metacarpal with small attachments to 2nd metacarpal mating season (Bernard et al. 1997). In a Malawian populationof and phalanx I.1 (Thewissen and Etnier 1995). Surface of pad is P. nanus, early spermatogenesis began in February and, by June, highly ridged with ridge surfaces appearing smooth when magni- late spermatogenesis was underway in almost all seminiferoustu- fied.Bulkofpadisdermaltissuewithlargesebaceousglandsthat bules, and prostate and seminal vesicles were starting to secrete areassociatedwithhairsandthatopenontoedgeofpad.Proximal fluids into lumina (n 5 18—Bernard et al. 1997). In June, thick- side of pad and side that faces radius have higher concentrations ness of seminiferous epithelium decreased, indicating no new di- of sebaceous glands. Stratum corneum layer of epidermis is thin vision of spermatogonia. By late September, spermatogenesis was and thickness of stratum malpighii varies but is usually thicker complete, but spermatozoa remained in cauda epididymides and than stratumcorneum.Elasticfibersareembeddedincollagenous secretory material remained in lumina of accessory glands (n 5 septaindeeperpartsofpad.Astrongconnectivetissuebandwith 24—Bernard et al. 1997). Reproductive males show distensionof anembeddedpieceofcartilageconnectsepidermisofpadonradial sacculum with elongation occurring during testicular hypertrophy side to adductor brevis muscle, which also inserts on metacarpo- andsubsequentfillingofepididymis,thereforemakingmalerepro- phalangeal joint (Thewissen and Etnier 1995). Because adductor ductivestatevisibleupon examination(O’Shea1980). brevis inserts in center of pad, it can raise the center forsuction. Prior to estrus, endometrium is thin (mean thickness 6 SD, PadsofP.nanusmaybeusedtogeneratefriction(LaValandLaVal inmm;97.568.1,n52),withfewshort,straightuterineglands 1977) or may function as suction pads (Allen 1939b; Smithers andovarieshouseprimaryandsecondaryfolliclesofmeandiameter 1983). Insertion of adductor brevismuscle allowssuction-cupfor- (6SD,in mm)127.4613.2,butnoGraafianfollicles(Bernardet mation,butrimcannoteasilybesealedduetosurfacerelief.Some al.1997).Byonsetofmating,endometriumisunchanged,butsec- deeper folds in center of pad may be sufficiently smooth to seal, ondaryfollicleshaveincreasedinsize.Ascopulationoccurs,large numbersof spermatozoacollectinupperendofuterinehornsand providing some mechanical adhesion, but overall surface relief of uterotubal junctions (Bernard et al. 1997). Female P. nanus store padsuggestsP.nanususesfrictionratherthanwetordryadhesion sperm(Bernardetal.1997).Vaginalplugshavenotbeenobserved (ThewissenandEtnier1995).SomeP.nanusspecimenshavesub- (Bernard et al. 1997). In a Malawian population, all femalescap- cutaneous fat in thumb pads (Schliemann and Schlosser 1978; tured in July had mated and had small Graafian follicles present Thewissen and Etnier 1995). Thumb pad is anatomically distinct for the 1st time (Bernard et al. 1997). By late July, endometrium from that of Glischropus and Tylonycteris and evolved indepen- wasthicker(187.7629.1SD,mm,n58),uterinewallwasmore dently(Thewissenand Etnier1995). vascularized, uterineglandswere longer and morecoiled,andav- Dentalformulaisi2/3,c1/1,p2/2,m3/3,total34(Rosevear erage diameter of Graafian follicles was 275.4 6 38.4 SD mm (n 1965). Upper incisors are nearly equal in length with inner ones 58—Bernardetal.1997). bifid (Dobson 1878) but not deeply cleft (Koopman1975).Degree P.nanusgivesbirthonceperyearatbeginningofwetseason of bifidy is highly variable (Rosevear 1965). Outer upper incisors (November) and lactates for 8 weeks in South Africa (Kwazulu- are unicuspidate and relatively long, ca. half the length of inner Natal, 288559S and 308249S—LaVal and LaVal 1977), Malawi upper incisor (Koopman 1975). Lower incisors are trifid and not (Zomba,158239S;358199E—HappoldandHappold1990a;Namad- crowded(Dobson1878).Anteriorupperpremolarissmallandusu- zi,158319S;358119E—HappoldandHappold1996;otherlocations ally just visible from outside of toothrow in gap between canine in Malawi—Happold et al. 1987; Lawrence and Loveridge 1953), and 2nd premolar (Rosevear 1965). Lower jaw is comparatively and Kenya (near Kibwezi, 28189S and 38879E—O’Shea 1980). In weak with no ascending ramus; a short broadened surface with a KenyaandMalawi,birthistotwins(HappoldandHappold1990b; low coronoid is present for attachment of jaw muscles, and lower Kershaw1922).Reproductionisaseasonalandtwinsareeitherrare part of mandibular symphysis is often strong, allowing for an ex- orunknowninDemocraticRepublicoftheCongo(Shabaprovince, traordinarily wide gape (ca. 1208—Lang and Chapin 1917a). Sag- 118399S;278289E—AnciauxdeFaveaux1983;58N–58S—Langand ittalcrestisabsent.Lamboidalcrestsarenotpronounced(Rosevear Chapin 1917a; Parc National de la Garamba, 38409N–48409N— 1965),andaconcavityoccurswhereforeheadmeetsrostrum(Koop- Verschuren 1957; Virunga [Albert] National Park, 1–28N—Ver- man 1975). schuren 1966), in Rwanda (1–38S—Anciaux de Faveaux 1983), Wingloading(6SD, n55)is8.462.0Nm22,meanaspect and in Gabon (ca. 18N—Brosset 1966). Polyoestry is not evident ratio ranges from 6.7 to 7.3 (Aldridge and Rautenbach 1987; (Happoldand Happold1990b). O’SheaandVaughan1980),andwingtipshapeindexis1.460.5 Asynchronous parturition occurs in Democratic Republic of (6SD, n 5 5—Aldridge and Rautenbach 1987). Maneuverability the Congo, Gabon, Rwanda, and Tanzania (Anciaux de Faveaux duringflightwasquantifiedfor6(sexnotspecified)P.nanususing 1973; Baagøe 1978; Brosset 1966; Lang and Chapin 1917a; Ver- strings suspended in a flight tunnel, and average minimum nego- schuren 1957, 1966); whether sperm storage occurs under these tiableinterstringdistancewas0.2060.07m(6SD—Aldridgeand conditionsisunknown(Bernardetal.1997).When84batsinthe Rautenbach1987). UsambaraMountainsofnorthernTanzaniawereexaminedbetween Baculum of Pipistrellus nanus is ca. 1.7 cm long. The ex- July 25 and August 7, no juveniles or scrotal males were found, panded base is divided into paired lobes and is sharply flexed to and only 1 of 49 females was pregnant (twin fetuses—Baagøe shaftwithdistalpartexpandedanddownwardlydirected.Baculum 1978).InGarambaNationalPark,DemocraticRepublicoftheCon- hasabulbousundividedtipinsameplaneasbase(HillandHar- go, females captured in April and May were neither pregnant nor rison1987; Kearneyetal.2002). lactating,andjuveniles(,1monthold)werecapturedinlateJuly Individualsroostinginleavesandinthatchedroofsusedaily (n 5 2—Verschuren 1957). In Parc National du Niokolo-Koba, torpor (Happold and Happold 1990a, 1996; O’Shea 1980). Al- Senegal (ca. 118N–128309N) females captured in May (n 5 9— though sustained torpor in cool weather is not possible given Verschuren 1982) were nonreproductive. In National Park W in ephemeral nature of furled leaves (Happold and Happold, 1996), southern Niger, 2 females captured in late February and early limited use of daily torpor occurs in a leaf-roosting population in March were nonreproductive and 8 males all had scrotal testes Malawi (Happold and Happold, 1990a, 1996). Some African ba- averaging3by2mm(Poche´1975).InRioMuni,EquatorialGuinea nanabatstakenfromfurledleavesweretorpidatendofDecember (ca. 1–28N), a lactating female was found in August and a scrotal (mid–wetseason)andallbatsweretorpidinJune(cooldryseason). male(testes5 by3 mm)inNovember(Jones1971). TorporwasnotobservedinKwazulu-Natal,SouthAfrica(LaValand In Malawi, the year can be divided into 3 seasons: hot dry LaVal1977). season(HDS)extendingfromendofAugusttomid-November;hot 4 MAMMALIANSPECIES 784—Pipistrellusnanus wet season (HWS) from mid-November to end of April; cool dry orifitmerelyusesitsclaws(Brosset1966)towedgeitselfintothe season (CDS) from May to August (Bernard et al. 1997; Happold tube-likeleaf. and Happold 1990a, 1996). Malesbecomenonreproductiveinthe Diet varies between individuals and season; small moths are HDS and females are pregnant. Fertilization success is close to eaten during cool, dry seasons, but seldom during wet seasons 100%(Bernardetal.1997).Parturitionoccursatthebeginningof (HappoldandHappold1996).Oneindividualinthedryseasonin the HWS, a period characterized by high night-timetemperatures Zimbabwe(188109S;288139E)hadeaten50%Coleopteraand50% andinsectabundance.Spermatogenesisbeginsduringthe2ndhalf Lepidoptera; in the same area in the wet season, fecalanalysis(n of the HWS and testes reach maximum size by end of April (in- 510,50pellets)revealed52%Coleoptera,40%Lepidoptera,and cludingthosemalesbornthepreviousNovember).Juvenilesreach 8%Diptera(Fentonetal.1977;FentonandThomas1980).Ability adult size by end of the HWS. Start of the CDS coincides with tocrawlmayallowP.nanustogatheratleastpartofitsinsectdiet release of spermatozoa into cauda epididymides, and mating pro- from within leaves and around fruit at its roosting location (Lang ceedsfrom mid-JunetoearlyJuly,whenharem-likegroupingsare andChapin1917b).Orthopterans,mainlykatydids,occurinfurled observed. In July and August, spermatogenesisand spermstorage bananaleaves,alongwithfrogsandlargehuntingspiders;noneof occurs, and further matings may occur. In late August, ovulation these shared a leaf with an African banana bat (LaVal and LaVal and implantation occurand gestationlastsca.10weeks(Happold 1977). and Happold 1996). Some Africanbananabatsathigheraltitudes Insomeareas,P.nanusroostsinthesamebunchofbananas ovulate later (Bernard et al. 1997). In Kenya, the chronology is as Myotis boccagei (Lang and Chapin 1917a). Interspecific com- similar,withtestesevidentattheendofMarchandmatingstarting petitionforroostsisunlikelyduetoabundanceofsuitablebanana mid-May (O’Shea 1980). Testes are present from April until Sep- leaves(HappoldandHappold1990a).InMasalani,Kenya(28189S; tember and gestation begins at the end of August with parturition 38879E),NeoromiciasomalicusandP.nanusco-exist(O’Sheaand inlateNovember.DatafromKwazulu-Natal,SouthAfrica,aresim- Vaughan 1980) and have similar horizontal gapes (measured as ilar,withtestesevidentfrombeginningofMarchtoendofAugust widthbetweenoutsidesoflastuppermolars;ratioofN.somalicus (LaVal and LaVal 1977). Pregnancy spans September–November, to P. nanus is 1.07), indicating slight to no resource partitioning parturitionoccursinlateNovember–earlyDecember(slightlylater on basis of prey size. P. nanus has a higher frequency range of than in Kenya and Malawi), andlactationoccursuntilearlyJanu- echolocation (sweep of 70–35 kHz for N. somalicus—O’Sheaand ary. Females with juveniles occur at all times of the year in East Vaughan 1980) and N. somalicus forages in more open areas Africa(58N–128S),suggestingaseasonalbreeding(Kingdon1974). (O’Sheaand Vaughan1980). Twinning occurs when 1 fetus develops in each uterine horn Sex ratiosvary with geographicallocation.InKwazulu-Natal, (HappoldandHappold1990a).Occurrenceoftwinningvarieswith South Africa, a 1:1 adult sex ratio (92 females:94 males) was geographic locality, ranging from 50% in South Africa (LaValand reported(LaValandLaVal1977).InZomba,Malawi,adultfemales LaVal1977)to100%inequatorialandsubequatorialregions(Hap- (n5141)outnumberedadultmales(n584)foratleast7months of the year, but equal sex ratios were observed for juveniles (n5 pold and Happold 1990a; O’Shea 1980). Lactating femaleswith1 or 2 young were obtained in March in Ghana (68109N—Jeffrey 58—Happold and Happold 1990a, 1996). In Namadzi, Malawi, 1975). In August and September in Zimbabwe (16–208S), most adult sex ratio varied with season and females (n 5 28) outnum- pregnantfemaleshadtwins(mean51.9fetuses;n526—Smithers beredmales(n512)duringHDS(lateAugustcensus),whilemales (n 5 24) outnumbered females (n 5 17) in mid-June (1st half of and Wilson 1979). Females with 2 young have been reported in Malawi (ca. 178S, 358E; n 5 3—Kershaw 1922) and Liberia (ca. the cool dry season), although evidence suggests that adult sex 4–88N;n57—Kuhn1962). ratios differ annually (Happold and Happold 1996). In Burundi (males:females, 19:1—Niort 1970), Democratic Republic of the Congo (277:85—Verschuren 1966), and Kenya (80:39—O’Shea ECOLOGY. African banana bats are widely distributed in 1980), adult males outnumbered adult females. Differential mor- rainforests,opencountry,andplantations,butavoidopenplacesof tality may contribute to skewed sex ratios (Happold and Happold villages. P. nanus often roosts in cracks and holes in buildings 1990a; O’Shea1980). Predation atroostsduringtimesoftorpidity (Rosevear 1965; Shortridge 1934), culverts, rafters, nearly full- may result in higher female mortality due to harem-like roosting grown bunches of green bananas, and dead leaves of musaceous arrangement (Happold and Happold 1990a). Predation by African plants (Kingdon 1974; Lang and Chapin 1917a; Rosevear 1965), false vampire bats (Cardioderma cor) has been observed and ar- palms(O’Shea1980;Rosevear1965),andpalm-leafthatchedroofs boreal snakes (especially Psammophis) may prey on P. nanus (Allen and Coolidge 1930; Lang and Chapin 1917b; LaVal and (O’Shea1980). LaVal1977;O’Shea1980;Rosevear1965).Individualsalsoroosted indesertednestsofWeaverbirds(Ploceus—HappoldandHappold BEHAVIOR.FlightofP.nanusisunsteady,nearlyfluttering, 1990a). P. nanus most commonly roosts in the long narrow tube withazigzagmotion,butnoabruptturns(LangandChapin1917a). formed by the unfolded terminal leaf of banana or plantainplants P. nanus is an aerial insectivore (O’Sheaand Vaughan 1980)that (Musa) and in curled leaves of Strelitzia caudata or S. nicolaii flies close to the ground (2.1–5.1 m aboveground; mean 5 3.6— trees (Baagøe 1978; Kingdon 1974; LaVal and LaVal 1977; Rose- Kingdon 1974) and feeds on insects within 1 m of vegetation vear 1965; Smithers 1983; Verschuren 1957), Ensete (Verschuren (O’Shea and Vaughan 1980). P. nanus flies continuously and, al- 1957); or wild loquat,Uapaca kirkiana(Happold etal.1987).In thoughitusesarangeofhabitats,spendsmuchofitsforagingtime furledbananaleaves,Africanbananabatsarelocated1–4mabove just above the canopy (Aldridge and Rautenbach 1987). Foraging ground (Happold and Happold 1990a), and plants taller than 1.5 under the canopy occurs (Fenton et al. 1977). P. nanus pursues marepreferred(Baagøe1978).Openingdiameterofthecylindrical smallinsectsaroundlightedlamps(LangandChapin1917a). tubeformedbytheunfurledbananaleafrangesfrom3to.24cm Foraging individuals have been followed using fluorescent (Baagøe1978;HappoldandHappold1990a).Africanbananabats powders and ultraviolet light (Fenton et al. 1977). Flanged metal arenotfoundinleaveswithopeningsizes,16cm2,andopenings bat bands have also been used to study behavior and such bands of101–950cm2arepreferred(Baagøe1978).Individualsroostfrom causedminimalinjuries(HappoldandHappold1998).P.nanusis 10–750mmfromtopofleafatpointsintubeaveraging38mmin activeonbrightmoonlitnightsandisthe1stspeciesseeninflight diameter (LaVal and LaVal 1977). Suitable banana leaves are not each evening. Emergence time afternauticalsunsetis43.164.5 limitingbecausemostplantshavenewleavesyearroundandbats SD min (n 5 118—O’Shea 1980). Individuals have a relatively occupy ,32% of suitable leaves (Happold and Happold 1990a, small activity range; recaptures occurred within 10–100 m of leaf 1996; LaVal and LaVal 1977). P. nanus prefers Musa to Ensete where they were banded (Happold and Happold 1996; LaVal and and Strelitzia (LaVal and LaVal 1977; Verschuren 1957), perhaps LaVal 1977). In a Zimbabwe colony, 3 short periods of foraging becauseMusaproducemoreleavesincloserproximity(LaValand activity occurred each night, with the 1staround 1900 h at emer- LaVal 1977). Furled leaves typically last 1–3 days (Happold and gence, the 2nd later in the night, and the 3rd just prior to dawn Happold 1990a). P. nanus has atypically been found in a cave (Fenton et al. 1977). Although movement is localized throughout (AellenandBrosset1968). much of the year in Kenya, a seasonalphaseofdispersalwasob- Thumb pads and somewhat flattened skull allow P. nanus to servedasproportionofdailysightingsofunbandedindividualsin- roostinmanysmallrefugesinadditiontofurledleaves(LaValand creasedandrecaptureratesofbandedindividualsdecreasedatthe LaVal1977).OpinionsdifferconcerningwhetherP.nanususesits beginningofthelongdryseason(O’Shea1980). padswhenmovinginafurledleaf(Rosevear1965;Smithers1983) Pipistrellus nanus roosts singly and in pairs, small groups, 784—Pipistrellusnanus MAMMALIANSPECIES 5 and colonies of up to 150 individuals (LaVal and LaVal 1977). and are described as ‘‘dee-dee-dee-dee-dee-dee-dip’’ (O’Shea Groupsizeandcompositionvaryseasonally(HappoldandHappold 1980:183).Thesecallsaregivenmostfrequentlybymalesinflight 1990a; LaVal and LaVal 1977; O’Shea 1980). During parturition whentheyarepassingroosts,emergingfromroosts,orencountering and lactation, adult males and adult females rarely roost together conspecificswhileforaging.Shortcallshaveeither3or4syllables (Happold and Happold 1996). Seasonal changes in social organi- and are described as ‘‘dee-dee-dip’’ or ‘‘dee-dee-dee-dip’’(O’Shea zationofMalawianbatsaredividedinto3periods:lactation,post- 1980:183).Shortcallsareoftenrepeatedmanytimesinsuccession lactation,andharemformation(HappoldandHappold1990a).Dur- and are given more frequently by males at rest in roosts than by ing lactation, adult males roost singly and lactating females roost males in flight. Short callsareoften exchanged betweenmalesfor with their young or with 1 or 2 other lactating females and their periodsupto20minutesandthese‘vocalduels’sometimesendin young.Duringpostlactation,adultmalescontinuetoroostindivid- anaerialchase.Vocalizationismorefrequentduringmatingseason ually while postlactating females (n 5 16) roost singly (19%) or (May–August)andinthelastfewhoursbeforedawn(O’Shea1980). withgroupsofsubadultsorotherpostlactatingfemales(81%).Dur- Theseaudiblevocalizationshavenotbeenreportedinotherstudies ing harem formation, testes of all males are scrotal and all males ofP. nanus(Happoldand Happold1996). and 95% of females(n538)roosted in groupsof2–9bats.Most groupsduringharemformationwerecomposedof1adultmalewith GENETICS.Pipistrellusnanushasadiploidnumberof36, 1 or more females; a few all-female groups were found, as were afundamentalnumberof50(PetersonandNagorsen1975),andis groups containing females with 2 males (Happold and Happold related karyotypically to Neoromicia capensis(Heller andVolleth 1990a). Composition of roosting groups is highly labile (Happold 1984). The GTG-banded karyotype includes 8 biarmed and 9 ac- and Happold 1996; LaVal and LaVal 1977; O’Shea 1980). Small rocentric autosomes, with a medium-sized metacentric X chromo- groupsof2–3individualsofunknownsexforagetogetherinsmall some and an acrocentric Y chromosome that is smaller than the areas(O’Shea1980). smallestautosome(Kearneyetal.2002).Inadditiontometacentric Malesroostwithmultiplefemalesandfemalesroostwithmul- chromosomes1/2,3/4,5/6,and16/17,4chromosomesresultfrom tiple males in a highly labile fashion during mating season, with Robertsonian fusions between chromosomearms 7/11, 8/9,10/12, no apparent male–female bonds (Happold and Happold 1990a, and13/14.P.nanussharesallbutthelatterfusionpairwithNeo- 1996; O’Shea 1980). Although the term harem has been used, romicia capensis, N. rendalli, and N. zuluensis (Kearney et al. groupsoffemalesarehighlylabileincompositionandthereforedo 2002). P. nanus from South Africa is monomorphic for the same not match the strict definition of harem (Happold and Happold allele in all but 1 of 13 liver-enzyme loci tested; 2 allelic forms 1996).ThematingsysteminaKenyanpopulationroostinginper- existfor glutamate dehydrogenase(Moralesetal.1991).Basedon manentthatch-roofstructureswasresourcedefense,inwhichmales protein electorphoresis, P. nanus shows some genetic similarity defendedroostsandforagingterritories(O’Shea1980).Inbothleaf- with Neoromica zuluensis and Pipistrellus hesperus, butdoesnot and thatch-roosting populations, females showed no roost or mate groupwithEptesicusfurinalis,E.fuscus,orE.hottentotus,which fidelity,whereasmaleswerefaithfultoparticularroosts(thatch)or grouptogether(Moralesetal.1991).P.nanusgroupswithP.pip- patchesoftrees(HappoldandHappold1996;O’Shea1980).Males istrellus,P.p.mediteraneus,P.mimus,P.stenopterus,Neoromicia roosting in permanent structures displayed the highest degree of somalicus, and N. tenuipinnis based on mitochondrial (mtDNA) roostfidelity(O’Shea1980).Suchfidelityisnotpossibleinbanana cytochrome b (Barratt et al. 1995). Using sequences from 12S leaves given their ephemeral nature, although P. nanus exhibits rRNA,tRNAVal,and16SrRNAmtDNA,P.nanusgroupswithLae- fidelity to plantations and clumps of banana plants (roostrange— photusnamibiensisandNeoromiciasomalicus,whogroupclosely Happold and Happold 1996); males in a Malawian plantation together,andwithN.brunneusandN.rendalli,whogroupclosely. showedgreaterfidelitytocertainclumpsoftreesthandidfemales. P.nanusdoesnotgroupwithEptesicus,includingE.furinalis,E. In leaf-roosting P. nanus in Malawi, despite the availability fuscus, and E. hottentotus, supporting the protein electrophoretic of suitable roost leaves, sex ratio data from roosts (n 5 125 bats) data, nor with other Pipistrellus, including P. abramus, P. coro- suggested males exclude other males from roosts, although mech- mandra, P. javanicus, P. nathusii, P. pipistrellus, and P. tenuis anism of exclusion was unknown (Happold and Happold 1996). (Hoofer and Van Den Bussche 2003), supporting its removalfrom Males(n550)showedfidelitytodistinctclumpsofbananaplants Pipistrellusintoa newgenus. with little overlap between roost ranges (13–28% of occupied clumps were used by .1 male—Happold and Happold 1996). REMARKS. The common name African banana bat derives fromhabitofroostinginfurledbananaleaves.Speciesnamenanus Malesinathatch-roostpopulationpatrolledinfrontoftheirroosts Peters,1852 isLatinfordwarf(Rosevear1965). at all times of the night throughout the year and aggressiveinter- Koopman(1994)recognized6subspecies,listingP.n.nanus actions between patrolling males increased in frequencyatnight’s end (n 5 116 chases—O’Shea 1980). Males also defended ellip- forsouthernAfrica.InredescribingP.n. australisRoberts,1913, soidalforagingterritorieswithalengthof24–25m(n52—O’Shea ajuniorhomonymofP.hespersusaustralisMiller,1897(Koopman 1966), as P. africanus meesteri, Kock (2001) revived Peterson’s 1980). Certain malesattractedmorefemalesthanothers(Happold (1987) declaration that the larger sized form from Transkei is a and Happold 1996; O’Shea 1980). In Malawi, mean number of distinctsubspecies.Twosubspecies,P.n.meesteriandP.n.nan- femalesfoundwitheachmalecaught5ormoretimeswithfemales was11.3 (n519males—HappoldandHappold1996),andmate us, are recognized for the southern African subregion (Bronner et al.2003). selection by females was not linked to roost-range quality; males (n 5 18) that roosted in the more reliable clumps did not neces- OriginallyallocatedtosubgenusPipistrellus(Koopman1975), Hill and Harrison (1987) moved P. nanus to subgenus Hypsugo sarilyattractmorefemales(HappoldandHappold1996).InKenya, males(n511)makingmoreaudiblevocalizationsroostedwiththe Kolenati,1856,basedonbacularcharacteristics.BecauseHoraćek and Hana´k (1985–1986) elevated Hypsugo to generic rank, the largestnumberoffemales(O’Shea1980). binomenHypsugonanushasbeenused(HooferandVanDenBus- Search-phaseecholocationcallsaresteepfrequency-modulat- sche 2003). Based on GTG-banded chromosome data, Kearney et edcallswiththefollowingcharacteristics:meanduration,4–5ms; al. (2002) transferred nanus to Neoromicia Roberts, 1926. Based meandominantfrequency(kHz),ca.70;meanminimumandmax- imum frequencies, ca. range 62–67 and 86–90, respectively(n5 on mtDNA sequence data, Hoofer and Van Den Bussche (2003) found polyphyly within Neoromicia, and proposed Neoromicia 5—KrugerNationalPark,SouthAfrica—AldridgeandRautenbach 1987;n53—Sengwa,Zimbabwe,FentonandBell1981;n54— nanus be transferred to an as yet unnamed genus along with N. brunneusandN. rendalli. Jozini Dam, Kwazulu-Natal, South Africa—Taylor 1999).Callfre- We thank M. B. Fenton for taking the specimen photograph quencies up to 115 kHz have been detected (Kibwezi, Kenya— andskullphotographs,theRoyalOntarioMuseumforaccesstothe O’Shea 1980). Intraspecific variation in search phase calls exists. skullmaterial,andJ.SkinnerforuseofP.nanusdistributionmap. 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