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MAMMALIAN SPECIES vo. 312, pp. 1-5, 3 fe Eidolon helvum. By Sheri L. DeFrees and Don E. Wilson Published 30 June 1988 by The American Society of Mammalogists Eidolon Rafinesque, 1815 Fidolon Rafinesque, 1815454. Type species Vespertlio eampyras helous Kerr by subsequent designation (Andersen, 1908), Peerocyon Peters, 1861:423. Type species Pterocyon paleaceus Peters, Leiponya: Jenink, 1881:60, Type species Leiponyx bitikofet Tentnk CONTEXT AND CONTENT. Onder Chiroptera, Suborder Megachicoptera, Family Preropoidee, Genas Eidolon, which in cues only the species E. helo, Eidolon heloum (Kerr, 1792) Straw-colored Fruit Bat Vespertilio vampyrum helows Kerr, 1792-91. Type locality not specie subsequently fixed as Senegal (Andersen, 1907) Peeropus stramineus. E'. Geofivoy StHisire, 1803:48. Type ‘ot specified subsequently fixed us Senaar by Koopman (191 Pterocyon paleacens Peters, 1801423. Type locality “Africa ‘restricted to Senaar by Matschie (1899). Peeropus mollipitosus H. Allen, 1861:159. Type locality “Gabon.” Preropus palmaran Heuglin, 1865:38. Type locality “Middle and ‘Coper White Nile and between Sensat and Fazogl along the [Bie Nie"; subsequently fixed ae Senaar by Keopman (1975) Peeropus dupreanus Schlegel and Pollen, 1867419. Type locality "EMadagasear” Nantharpyia leucomelas Fitzinger, 1866:544, Type locality “Up- ‘ubuequently xed a Senaar by Koopman Leiponyex buttikferiJentink, 1881:60, Type locality “Liberia, St Paul's River (Millbare) Preroeyon sabaeus Andersen, 19073508. ‘Aden” (Saudi Arabia Eidolon heloum: Andersen, CONTEXT AND CONTENT, Context same as for genus ‘Three subspecies are recognized (Hayinan and Hill 1971) Eh, heloum Kerr, 1792:91, see above (stramineus E’. Geoffroy StcHlsice, paleaceus Peters, moliplosus H. Alen, pamarum Hugin, oncomelas Fivinger, and butikoferi Jenin are sy nym). Eh. dupreanum Schlegel ane B. k sabacum Andersen, 1907: Type locality “Labs 1912.2, First use of current name len, 1866:419, see above, DIAGNOSIS. Dobson (1878) united Bidofon with Rowsetus in the enue Cynonyctris, but Andersen (1912) demonstrated their tistinet generic status. Eidolon has a larger pl and a relatively longer rostrum, character sttes that might be considered more primitive than those af Rousertu, the genus with which it most ‘losely related (Andersen, 1912). However, the development of short, bony auditory meatus dintintly separated premexilarics fin contact oF often fused in Rouseurus}, a lengihened MI, a reduced M2, clearly vsble sexual differentiation: in color (abeent in Rom seitus) and generally larger dimensions suggest that Eidolon more Specialized. Eidolon has a greater wing span than Rousettu, its wings are more tapered and pointed, and its Sight is more linear (Kingdon, 1974), The second digit i clawed as in Rousettus and Preropus. Wing membranes extend from the sides of the dorsam and the back ofthe fst toe in Eidolon and Ptoropus. Eidolon has ‘til Tes than half the length of the hind foot with 2.0 to 2.5 rebrae protruding (similar to. Rowsettus, slightly longer than Myonyrters, Pteropus has no tail. In Eidolon, asin Pteopus, the busiranial axis defected; however, the occiput sneither elongated nor tubular. The palate broadens posteriorly, is widest between M2 to M2, shen narrows a the posterior border toa with approximately tual fo that between the lingual edges of P4 to PS, This dimension istmuch narrower in Pteropus. The length ofthe rostrum is mueh {greater than the lachrymal width ofthe sellin Eidolon as compared to Preropus. In Myonycteris, the [ront ofthe orbit i located ver Teally above the anterior portion of ML, whereas in Bidolom itis located above the middle to posterior portion of MI (Fig. I) GENERAL CHARACTERS. Short pelage covers the head, dorsum, and venter ofthe strav-coloed frat bat. On the neck the fr is thick, and nearly absent on the face in font of and below the eyes (Fig. 2), The ears are hairless posteriorly with far at thee base (Andersen, 1912). Pelage extends Over the upper arm and slighty fonto the forearm and upper surface of legs at interfemoral mem: brane, but not onto the wing membrane (Nowe and Paradiso, 1983; Fic. 1. Dorsal, ventral, and lateral views of cranium and lateral view of mandible of female Eidolon helvum, U.S. National Museum of Natural History 411810, from Ghana. Greatest length of eal 53.6 mm, Fic. 2. Eidolon heloum photographed in Kenya by Merin Tut Smithers, 1983). Despite its name, the color of heloum isnot a ‘consistent shade of golden-yellow, Color vaties between areas of the body ofthe same individual as well as between individuals, However, the general appearance ranges from a pale yellow-gray to 8 dark sepaogray- Adults usually havea bright orange to deep tawny-colored collar whichis brighter and more pronounced in miles (Rosevesr, 1965). The young are simlar in color to adult males, although severally darker and lacking a collar (Andersen, 1912). Individual hairs are buf atthe base and tipped with brown. Hairs on the venter ace yellovih-bu Iaterally and brow in ealor midventlly, Males appear darker due tothe longer brown tis of the baron the bead, lower part of the buck, and dorsel sie ofthe tibiae. The bar of the calla ie longer than the remainder ofthe shor, fine, closed fur ‘Wings of this species are long, pointed and romewbat narrom and are a dark Hlackshdrown, At rest, the ends of the wings are folded back wit the tip folded in; the second phalag and fourth digits be lat against the lower surface ofthe wing (Allen fetal, 1917). Wings extend from the sides of the dorsum ease to spine, and from the back ofthe fist toe. The metacarpal and frst phalanx of the fist digit bie within the membrane (Andersen, 1912). Wing areas average 739.4 em? in males and 545.3 em’ in females (ones, 1972). The upper surface of the tongue contains pattern of diferent kinds of paplle, The central pepllae, which can be seen withthe unaided eye, are rape and direetod posterioriy. The thumb is Tong and terminates in a sckle-shaped elaw. The toes alo have siekleshaped claws (Reseveas, 1965) ‘Serual dimorphism in sizeof E. heloum isnot great (Andersen, 1912). lenes (1972) found mearurements of forearms of males to average 19% greater then thove of females, Ehelaum is the second largest ofthe West Africen frat bats, exceeded in size only by the hammer headed frst bat, Hypsignathus monstroas (Okon, 1975 Rosoveaz, 1965). Ranges of external end cranial measurements (in ‘aun arr total length, 150.0 to 195.0; length of forearm, 117.0 to 182.0; wingspan, 750.0 to 950.0; length of thumb, 42.0 to 50.5 digit length of metacarpal, 51.2 t0 62.0; length of phalanx I, 15.3 to 18.5 length of phalanges IL and Ull, 10:5 to 16.8: digit Tl—eagth of metacarpal, 76.5 to 88.8; length of phalanx I, 50-2 to 56.2; lengths of phalane Il, 74. to 91.5; digit [V—length of retacarpal, 74.0 1086.0; length of phalanx I, 39.8 to 46.8 engl of phalane Il, 47.2 to 55,0: digit V-—length of metacarpal, 70.5 to BES; length of phalenx I, 30.0 to 39.7; length of phelans Il, 32.310 :40.0¢ length of ear from nore, 27.2 to 28.0: greatest with of ear, Nattened, 18.5 to 20.0; font of eye to tip of mule, 23.2 to 25.0; length of tal, 10.0 to 15.5: longth of tibia, $7.0 to 52.0; lengths of hindfoot, 31.5 to 38.5; skull—total length to front of premanilries, 54.5 to 62.2: width of brain eae at 2ygomata, 20.8 to 23.5; sygomatie width, 32.0 to 36.0: width between M2s, ex: ternal, 16.0 to 18.0; width across canines, 9.8 to 11.0 posterior palete to incisive foramina, 26.0 t0 80.0; front of orbit to tip of hasals, 19.0 to 22,8; length of mandible, 43,0 10 49.5; length of ‘mauilary toothrow, 21.0 to 23.8: length of mandibular ootheow, 28.0 to 26.2 (Andersen, 1912; Fayenuwo and Halstead, 1974; Kingdon, 1974: Rosevear, 1965). MAMMALIAN SPECIES 212 Fic.3. Distribution of Eidolon helewm, Central area is occupied yearsound, and migratory routes extend range north and south, ‘The continental subspecies (1) is. heloum heloum. The Arabian perinwula form (2) is E. . sabacum, and the Malagasy eubspecies (3) Bk dupreanum. The dental formula of E, heloum i i 2/2, ¢ L/L, p 3/3, m 2/3, total 34 (Rosevear, 1965) Teeth are without special medi fiestions and no secondary cusps appear in eanines or cheekteeth Molazs contain » longitudinal median groove separating» higher ‘outer and lower inner ridge. The upper inciaore are small, rounde ‘nd suboqual in size, Lower incisors are simular to uppers and wrually in contact with exch other and with canines, All postcanine teeth ste slightly separated. Ten palatal ridges are presets four anterior, three middle, and three posterior (Andersen, 1912). ‘Kingdon (1974 gave the range in body mare of F. elewm as 250 to 311 g. According to Fayenuno and Halstead (1973) the ‘mean body mas of males during the breeding season increased fom 230 ta 330_g and the corresponding mean body mase of females {rom 240 wo 350 g In Females the ineease was due to the developing embryo. DISTRIBUTION. Anderson (1907) reported the AVricen Aistibution of E.heloum to be from Somaliland (Somalia, Dut, Southeastern Ethiopie), Seneaar Sudan), and Senegambia (Seney ‘nd Mal) in the north, to Nyasaland (Malaw), Namaqualand South ‘west Africa, South Alriea), and Mashonaland (Zimbabwe) inthe uth, The straw-colored fut bat isa migrant from ts prime habitat, the tropical forests of the central part ofthe African continent, which supply the bats with a variety of fuits throughout most of the year, {to areas both north and south of this optimal teetory (Fig. 3). The swalabity of food determines the occurrence of this species cle svhere. E.heleur hos been recorded in forested habitat rom Guines to Nigeria and Cameroon, Galo, ard Zaire to the Rit Valley ard parts ofthe Sodan, Uganda, Kenya, Tanzania, and northern Angola Kreas where E. heloum miay be considered @ year-round resent include parts of northeastern Zambia, Malawi, and possibly north astern Mozambique and nortbeastern Natal (Sithers, 1983). Hay- na and Hill(1971) reported an even wider distribution from Senegal to Sudan, Ethiopia, Tanzania (including Zanzibar, Pemba, and Mafia Islands), Zimbebwe, and South Africa. In the west, the range covers the islands of Fernando Poo, Soo Tome, Principe, and. Ansobon. Outside of the African continent, these bts are known from the southwestern Arabian Peninsula and Madagascar (Fig. 3). Records fre infrequent from Namibia in Zimbabwe as far wet as the Maropo Hill, and in Mozambique south ofthe Zamna! River inthe cental snd southern parts of the country. In the southwest provinee of Transvaal these bate are considered “wanderers.” In edition, E: Aeloum bas been taken in the Orange Free State and inthe eastern Cape Province as far west as the Bredasdorp distri (Smithers, 1043). Rosevear (1968) reported that this species had been taken cout at sea 250 km from the nearest land, It has been recorded an ral islands off the Cameroon (Ambas end Biba) and Sierra MAMMALIAN SPECIES 312 Leone coasts, and on islands in the Congo Ri Laie). “There is no foeil record for thie specie, and in Lak Kira FORM AND FUNCTION. Te skin underlying he bighter colored calla behind the head te composed of an epidermal layer {ve to three cell tick in addon to «dermal layer containing merous sebaceous land. Those glands are better developed i Males Gann fmalesand arent essen injvendes(ainoye aod wel, 1979; Mutere, 1967), In mals, glandular yer compose ‘upto 79% ofthe total kin tikes in feralen up 1 42% (Malays Sid Hoel, 1979), These glands scete 2 muyaeling fui oe rolucton of whch hasbeen obecrved to nceata when nda ‘lsturbed (Alen et ab 1917 Kingdon, 1974). The ippes are tilay (Resear, 1965). ‘Eidolon Reloum hax ben wed in ties of the development of the web manln of the wing sd the org of stl eal ‘According to Church (1969:480) "The web arses as a bud on the itera by wal ps need sarky hy axon. The we mien ferent; paternal dstdstionpresgig the sla. ‘Hgement.* Before bir, cells remain dived inte two types nul ers and surrounding satelite cl The Inter supe myo- tiotnJovehyng ae regenerating mare and are ost teers the basrent and plasma meneame of mus bere (Muir eta, 1968) Laan and This (1968) sed single fbr ol motor terves from web mics. The mean lg of 57 inated inact fiers was 141mm. The endplate loci he within the mile alt each fier, generlly ear the center ts aw the heart of tales is sgnifeantly larger ereloped han that of female ial respects xcept forthe thickness of the right venricuar wall. The anterior ‘enn cavainiense compared tothe poster sana cov reflecting the irultory system requirement for fight. The free margin of the sins septum forme an exceptionally thick, masculr rg be- ten the lt anterior aud posterior vena cava (owt, 1967) During fight the anantmeces clu, daconnectng te aeres snd veine ad easing the Hod to Bow trough peripheral xpllares inthe libs, which are bypased le the bat at rest. The bear receives a sulden icreaae in vets ood. & pres for whic it Tas been moved In both mules and females the wal ofthe cons Gr oto tract ofthe right veil, conadrably thinner than that ofthe ins or flow teat. Th the tremendous ineesre ‘enon return esc hy fig cb accurate hy the ert {Kingdon 1974 Row, 1967). Tiood and bone marrow cele ofE. helm were extnined for respons o Skin back B and peril acitSchif postive mater, Peridee napibel ASD chorenctetsentease, at bela gucuror Tite. Strong perordae rencinty vas seen ‘only in developing iamloeyes and tmonoeyer of bone marrow. nese peroxidase Ractny was observed in developing and matre eryvocytes Hid and one marr (Cantona, 1977 ood pasage in E eloum ie rap Ronevear, 1965). Rodin ann Bequaert (1916) reared thse bats in captivity ona dc of set inane This frit pated through the alimentary canal wih short porind of time’ and war pated cut easy unchanged except {er the jes, The sophaguehapprezimately 100 mm lng and is marked by two datnetanatontel features uncommon song tmarunaly a apneic prtrotn athe gastocsopbagealjuneton Compored of mer eelar msde and 4 wel develo Inucone atthe base of the exphegts occupying approxin ‘nm of the cophagis sores incon wth the stomach, The Somach, (ally dstended, measures approtnaiey 90m and com St of four tegions—fundus, corp pyloric antrum, and pyre anal, Each wcton i larly marked, the fn beng nalieesy trier than the corp an the pyloric mh narrower (aout an indianerer A pect feature ofthe stomach i the presence of dep, pertanet furrows inthe ontreurace of the pyoic antrum, ‘The presence of both chief and parietal cells nthe Pyorc mca of the stomach i unusal abo, pining tothe posbity that. ht have poets carnivore or might reflect eeding er type of proeinrich food source, sacha pen, No topographical difference exe between the large ad sll iesines ievel they are dstingusable atthe hielo! level. Inthe Protinal dvolenum there i atanationl rego said tobe ich a floretype lands and in Brunner’ glands Urano in (rt bats Banners glands might seve to protect the code rom ler: ation by wsipepan, One sie of the sal inten theker than 8 ‘he other: the thn half is rudimentary, whereas the thicker is fully developed. The large intestine i measired as the final 30 em ofthe ‘entie length of sina and large intestine, which together measured bout 170 ex. Absorptive eels in the lage intestine are abundant, ‘wheveas goblet cells are not. The appendix and caecum are lacking 4s in other bats (Oken, 197), ‘The presence of enzyme-producing glands along much of the length of the alimentary canal contbutes to the feeding efiieney of these bats. Trypsin, pepsin, and amylase are produced ever in the distal esophagous invertase and maltase are found everywhere ‘except in the eophagus and stomach. These enaymes aze used in ‘earbebydrate and protain digestion, which also suggests carnivorous food habits, although they may be used to digest pollen, Peptidase, lipase lactase, and cellulase wee not detected in E.heloume(Opunbiyi and Okon, 1976). Despite considerable diference in pH values Fequited for maximal activity, pepsin and amylase are found in al four regions ofthe stomach, suggesting «regulatory mechanism for controling seretion events (Okon and Ogunbiy, 1979). Intercapular brown adipose tue undergoes marked diel ‘changes. More smultloealer cals appear in the afternoon: more tunlocular cells inthe evening, and they increase in number during Hight. A high nuaber of multioeulate cells inlestes high metabolic activity. Thus, brown adipose tissue contributes to daily energy ‘exchanges in ths species (Okon, 1980) “The body temperature of E.helcum has been recorded from 30 to 40°C; usualy from 32 19 37°C Jones, 1972). The uterus is Iieoenuate Both horns and thei corresponding ovaries ar functional (Mutere, 1967). ONTOGENY AND REPRODUCTION. Breoting is ses onal, with most copulation occurring from Apri to dune (Mutere, 1968). The ogg is fertied and develops unt the blastocyst stage but docs not continue development antl implantation in October (unmilayo, 1979; Mutere, 19650), Births ake place fom February {o May prior tothe onset ofthe higher of the two rainfall peak ‘The minimum gestation period is about nine months, although true station lasts only four months. Females produce one infant Soncy an Hants occurs in ateray flees ta are este a females (Funmilayo, 1979: Kingdon, 1974; Mutere, 1965b, 1967, 1980), ‘Mean weight of testes increased from 1.0 to 2.5 g (October Noveilct) to 5.2 to 5.5 g (April-June an increase caused by the increased nurer of spermatona inthe epiddynsdes(Mitere, 1967) Tones (1971) listed average dimensions of testes (in mim) a3 20 by 1 and 21 by 16 (May and June). Ogivie andl Olive (1964) gave testicular dimensions of one specimen a 15 by 12. Spermatoeoe can be found in the female genital tract from April w June. Ex: mination of uteri reveals unimplanted. Hasty from iy to September. Reasons for delayed implantation remain obscure (Mu tere, 19654, Pregnancies occur in either the right or left horn of the uterus never bh; «functional corpus lateum corresponds to the side of the hor bearing the embryo (Muere, 1907) Te fetus has it ibs crowed and tucked near the body with wings on ether side of and covering the face, ‘The head f large Compared tothe rest of the body. Fayenuwo and Halstead (1974) fecorded an increase in fetal weight fom 1 to 5 g within 16 weeks. The uropatagial muscle becomes well developed before bir. The yes of newborn E. eleum remain closed for several weeks. Hind Fins are well developed for clinging to the mother. Infant, which suckle upsiledown with one foot holing on across the mother’ Ttomach, are often attached toa nipple ofthe mother. Milk. teeth are present at birth (Halbtead and. Middleton, 1975). Consinual rooming and attention from the mother ae eseential forthe survival Sf newborns (Halstead, 1975). Joveles averaging 120 10 150 Sow separation behavior from mothers hy maldne (Funmiayo, 1979). Young males are not sexually mature by the folowing mating season (Halstead and Midleton, 1975). No spermatogenesis occurs uns testes reach a mass of 0:9 g (ata body mass of 240 to 249 1). Females cannot reproduce unl they have attained a bay mass 1 200g. No record of femalosin breeding condition exit in southern ‘Mica suggesting the improbabity of breeding there due to low population density of thene migratory animals (Mutere, 1967). The ‘ely known lngevty record far E.Reloum is 21 years, 10 monthe (Nowak snd Paradiso, 1983). ECOLOGY. Ridolon helvum feeds entirely on friting and flowering trees (Wilson, 1973) and in turn is peeved on by snakes and carnivorous birds and mammals (Funmileyo, 1979). Roost sites