Chapter 302 Zebra Reproduction Cassandra M.V, Nufiez, Cheryl S. Asa and Daniel |. Rubenstein Introduction Fw studios have ineetigatd the reproductive biol: ogy of wild equids; mach of what te know about bra specie hs come fm discon flowing calling Zoo research has provan usefal in deter mining vital details regaring the ovulatory cy- cling of females* ane the timing of puberty ard breetng life in both inales and females. Beha fora sttos both in th fled an in eapiity have helpet to arity these factors In adition, behav~ foral studies have reves! hove breeding behavior, the degree of breeding vensonality stages of pat sition, motherinfnt behavior, an the clispetsat of ‘Young males anu ferales ean vary inthe diferent Species Usirg this combieation of behavior, pPhysological and demographic dat, we explain the reproductive biology of each age and sox clas of pins sera (Equus burl, the tro subspecies of ‘mountain 2ebza ~ Cape mountain 2obra (Epis 2 va aera) and Hartmann’ zebea (Es zebra har nana) ~ ae Grevy'saebra (Eqs grey) ante Jats zelevance to ther eclegy al natal Nistor Social context: mating patterns and their ecological determinants Zebra live in two broadly ferent types of sox ties In one, pili by plains and mountain 2=- bra, females and mals ive xmimaleanalifemate family groups with thei infants, yearlings, and a= Yyearolss. These so-called ‘harem’ gioups ave ree wo sable eth anaes and females living togeler for any years i the other typified by Grey 520 ras, female associations are shorted and fens often roam in groups without males. Fasing and Ussing of individuals creates these open ener ership groups that wander through territories of ‘Goat eval. Prom the perspective of males, bath systems are polygynous since repredactively active tnd domisant males oon mate with more than one female. In pins zebras, polygyay involves hater defenge since females form tong associations, while in Grevy's aebas it nvolune resource ceense as males deiend grazing areas that females seek, As in any paygynous system, younger, less dominant rates form bacheloe groupe and attempt fo cuckold ions while stengitening testo females. Prom the perspictive of females, however, these mating systams clice diferent reproductive oppovtites In plains sabes, fomales are monaandous since sll females ia a harem bond to a paricslar male. In Grovy’s2b205, females ae polyandrous since they ten more actors lortorial Boundarae and mate witha vary f males in succession, Bach mating system emerges a8 a response to particslar scologial ctewmnslances.™ In general, Zebra mals provide lmporant material zeWards 40 females tat increase thir reproductive sucess. BY incrensing vigianee and enabling females 19 move ‘unhindered when seeking resources, some males tenable feral to spend more time grazing. Fe rales asceisting with the most daiinant males, of thee ising eapaly in rank, rae the mos offspring to the age of independence But ths only applies Jn mesic Habitats where food and water ~ the fa ey resources females need sustain themselves and their young ~ ace loeated lose enought each fther so tat Females of cifeent reproductive sates fan stay together and simltancounly derive these tume-elatatbenalits hat males provide. For Grewy's obras iniablting more al areas where aia is highly unprediciale, food and water are typically fa apart, us females having diferent needs cannot stay togetter an the social fabric becones torn. AS ‘result, fenae associations are weak. Since females exhibiting ether postpartum or eyling estrus are equally Valuable to males, those of high rank defend teritoros along routes to woter so that they can ‘mate wit atating females residing neat water andl "jen Resin Second Bon Bae by Anger O Nomen, a Sui Wel E Vn Diao Var ‘2500 Bice Pbshang Li2852 Mare: Other Equids norlactating females that must come to drink every 355 days’ Males of lesser rank defend teritores in grazing ares and thas gain access to 9. mote Iimited pool of rproductivly active females tis within these social and ecological contexts that thoreproductive biology of ebras mut be examine. Only by appreciating how socioeclogial selective pressures shape socal relationships and mating be- [havior can the silaciies and differences in opr duction exhibited by the varios age andl sex clases ‘ofeach zebra species be understood Mare Behavior Female female bonds in all ara epcies are weak ‘Theres litle compeniion over food and cooperative interactions in terms of llorooming o mata de- fense of young are rare. Sinaly, there i minimal competion for mates, bat female mountain zbeas compete for dominance, ad zank afletsreprodc- tive fess. Although both damiean! and suberd- rat famaloeprodaco 2pprocinatly the sare am Deco tispring over lifetime, the survival young, ‘born to high-ranking females exceeds that of young ‘baum olow-ranking feral” Insdie, a8 dang tees take on the rank of thei ethers, dominant f- ‘males will produce disproportionately move female ollsping. nal species, bond betiveen mothers and their young weaken as offpring mature, with bot males and females leaving their natal groups by 3 yea of age. Reproicing ferles and males inter ‘ct regularly, Females goreallyiatate movements, ‘withmnles directing the group fom the side or the rear Lactating ferales often its moves to water And be the way ‘Miing often involves confit among the sexes Females typically adopt a rigid psiton wit ta nice when iiiting copulation. More often, how ver the mae iene females in heat by sing or ldking the genital region. Resting his Head on Fenne’scumpor biting athe withers often precedes rmountighy hema Ira female isnt reat mate ‘he ether walks out fom under the male or uses her Inalegto drive him away Seasonality rebra In general zebra ar es seasonal than other Afican species. Plaine zebra foals are born throughout the yest although peaks do exist uring the lng eins Gigire 23) Kling veporet that BSS% of ole sere born from Octet Mare in he Ngorongoro Crater, Tanzani.Similany, Smuts found that 82.5% fouls were born between Saptenber at Ape in Kruger National Prk, South Aiea. Since the gesto- tion feriodis about 12 month, mating activity mast ‘occur tbughout the year with a penk during the ‘Sumner months Plains brain North American and SE Nig) Gop Os Now Dee in Fob Mar Ap Mey an Month ign 20.3 Sexton arto ith ore dierent speci ot compe me apt rom Kge (ines) oats (Cape moana ara esse Aare al Haines zt} ond Daten al? (rey aba ey en Seton copie animaI Fane setias [cues cts | | Al il | Zebra Reproduction 2853, ‘gue 022 Sera tution ith fr a ee acl Gey ih hein oa ts rae a te Pa aba uropean 2008 have given bitth in every month ofthe yeas, but Bicths lister in epring and eum ‘mer (Bure 3022) However trends for sping at Summer bits in zoos are sometimes the seit of ‘management decisions for young tobe bor in those month Mountain zebra Frceranging Cape mountain aebra mazes in the Mountain Zebra National Park at De Hoop Nature Reserve, South Attica fal year oun with peaks around the rains. Over 68% of Cape mountain fosls ae born during the rains, from November through April? Preeranging Hartman’ zebra in southwest ‘Aiea are similar, with 81-86% of fooling oecurrng from November through April? During a 3-year sudy of Hactmann’s zebra, mating sctivty was only Seen from September to April with a peal in Feb ry Phat data, coupled withthe at tat the get. perio i abot 1 yen inca tha ning ei inthe rest of he years quite rave. Grevy's zebra Grevy’s zebra are perhaps the mest seasonal ofthe zebras with prs of mating occuring during the log rains Fly and Augist and the short rns October and Novembor® Inthe fe ranging pops: lations of sosthern Kenya, breetng is highly depen dent upon avilable resources, with pas in ess oinclding uth higher resource avallailiy Con vers, mares can be induced ito anestus when resounges are setca® Gestation i bit longer i Grevss zabra (about 380 days). so peaks of rat ing setvity mast cecur in June theough Jay ant September hroagh October Captive animal in Dvr Kralove Zoo in the former Cuechosovakia gave bith every moath from 1973 1 1985, altough peaks did ‘exist in aruary and fom Jane hough Agel ges zal following the falng schedule of will popula tions Gee Figure 202.1), Data fom the Gevy's Zebra Iatermtinal Stadboole forthe petiod 198 trough 2004 show simular tend, sith births throughout the year but concentrate! more in spring and sum mer months. Av with plain zebra, this tend inay Feflect management strategies, but yearround en Aecriae monitoring ofthe her of Grevy’s rb at the Sant Lou's Zo0 (LE. Bauman, CS. Ast, and M. Fiche; unpub) has evened a pattem of lower concerrations of progestecone flor ovulations in winter, suggesting that ferlty might indeed be lower sing tht peso Puberty and breeding life Paine zebro Barly work on plains ebea defined puberty in nares ‘asthe onset of first eats and errous behaviors King reported fis tus to nocur at an average age of 2yeats4 months, ranging rom I year 8 months 2 years 9 months. Klingel® found thse Ngorongoro ares exhibited list ets ais, at 19-15 months of age Sirlrly, Smuts found thet the ist ests of2854 Mare: Other Equids “nkle321 Repro dla fer pins ond Grey's fede Pains ebro seat Nimberefenss a4 0 Ywongeiogheaest i yruimonts —2yeas6 rane Slastogyetiih——2eyans Tow 2Bye mere ox oes ‘soe ne te Nonerofms a8 ® Teoraeiowe fig Fant Ty 1 te Oiesiosectizgh—I6yees9 nenhs 2ye39 monks “ae ag stn men Bieceetemaieen Selatan san se aie SSO ane smaresin Kruger National Pak genre at] Sto 2yeaes ff age. The Pains Zebra North American Regional Siuook (ible 302.1)” has paeuetion records lor females at I year and 8 months! year and 10 months, land 1 yes and TT orth of age the data represent the femae’s age atthe time she gave bith, not the time she conceived. Klingl” estimated the reproductive potential of mares as oe cal pr yes fom the age of 50 about 15 to TB years They chsecved 120)mares forthe smnsors. Fitocn percent of the animals had thee foals (one pot yen), 3% hl wo, 42% ad one, and 10% had no offspring. On average, mares had #8 viving feal once every 2 yen. This coincides with ta fom a exptive ania at Chester Zoo, England ‘hat at 17 years oe was sl breading after having produced sx foals since attaining sexual matty? North Amevican Regional Seadbook (Table 302.1) recom show tha 10 females gver 20 yee of age pro- duced foals, the oldest at 26 years. Twelve females ‘gave birth to more than 10 foals over a ietime, OF 1232 recorded bets, 1 (8) were tring ‘Mountain zebra Jn Mountain Zebra National Pack, young Cape mountain zabea mares leave their natal hes af a ‘mean age ol 1 year 10 months; often they aren ist estrus this te’ Loyd and Rasa” reported Cape ‘mountain ilies in Be Hoop Nature Reserv leaving abit eats at a mean age of | year 7 mont (agin Aturing fist estus). Fe Foaling has been recorded at ages 25 young as 2 yeas!” to a old 485 years 7 mots” Regarlles of the exact age at fst fling, itseoms Hkely that the mares do na becoitesexally ‘mafurountl several month alter fst ests, The tual age of fist eats in Hartmann’ zebra hes pot tbe carded, bus fee xanging mares have thee st foal at about2yearsof age Young mares are expelled for the natal group at 14 to 16 months of age” at Which time they may be expevencing fist estrus. ‘The reprevctive potential of Cape mountain 2- ‘bra mares hasbeen estimated a about one foal every 25 montis, with ange of 13-69 mors? Sint lany, Loyd and Rasa” calculated an average inte bist interval of 27 months. Given » reproductive Mtespan of about 15 yeas, a Cape mountain zebra mare ean be expected to produce eight to nie fale inher lifetime. The epeedctive potential of Hast mann’ zabra has sot boon reported expliily, but may be surmised by maternal behavior Hartmana’s zebra mares wean their (cls at approximately 10 months of age, and begin to actively expel the foals 113 fo 16 months of age, shortly before the bith of hermes fol, Given this bebavioy, we can expect the reproductive potential of Hartman's zebra mares to bheabout one foal every 10 15 years, Grevy's zebea Palaty a fearing Grow’ zebeo mates has ber ‘estimated to oer at 3 tod years of age! with mating foccurting during the secon ests and conception feccurting after two othe estows cycles, However ‘captive animals have been ceported t reach puberty ana achieve conception at mach younger ages. The Inormational Subock as record of one female at? years 6 months an! another at 2 years IO nth gis- ng bint forthe frst ime (able 302.1) ta a study at the Saint Louis Zao tht maniowed behavior and fecal hormenes* a 2yearold female conceived but borted an 18m fetus at about dy. 100 of gest ton. She subsoquently mated arel aborted at lest ‘one snore tine before succesfully carrying afl to tern, which wasboen when she was3 yeas b months lof age. Managers hn suspected tat young females were more likely to experience abortions and the re Sus ofthis sdy supported that hypothesis. This “suggests that females may beable to vue and cone cave booze thir proactive tents ae slently ‘mature to maintain pregnancy. Age at puberty af [ected by ntrtion, but may also be intuenced by 50 ial contents in the wld that may nol be presen a captivity Free ranging Grevy’s zebra mares nay bree frm, 1 to few months afer fang, with am average re productive potential of one foal every 2 years!" Jn captive animals, the meen interinh hitev Inas been estimated at 18 months (ange 142-318 mont”) and 9.7 months ange 13-27 months) in (Czechoslovakian and Nout American r00, respec: tively. Howeves, management strategies likely binsthe average intervals, According othe Internationa ‘Stuabook (Table 3021)2 the oldest female to give thavas 28 years age, with 10 females over the age of 20 giving bith. Over» ifetine, there were seven females that each prodsced more than 10 foals, one ‘of which gave beth 013, The feral reproductive organs Plains zebra ‘The female reproductive organs of culled animals vere studied in detail by King The plane zebra mare has fwo ovaries, ich about Sem in lengthy ‘The ovaries cannet to the fllopian tubes (vid), ‘which conduct the ovum fram the ovary to the ‘wer, Each oviduetataches toon ofthe oo hoes of the uterus, which are approximately 18cm in length. The homs attach to the body of the uterus (00-15 em long). Caudal tothe ute the cervix, bout Sem long, wich ener the vagina, The vagina fs about 15-20cm long, and i quite elastic, allowing ito expand. The external opening of the vagina les ventral tothe rectum, The mate has vo manny lan, which are smaller in virgin mes. They ave To ducts each, hich open externally, Mex Detaled dissections ofthe female reproductive or ans have nol been reported in anountsin zebra Given their average macs of 234g and 276g for Cape mountain and Hartman’ eebea mares respectively they are likely to have organs ite termediate in size and shape to those of plains and Grovy’s zebra. For example, Westin van-Aarde {tl reported thatthe ovaries of Hatmana’s zebra females are heavier than those of plains zebra at ev ‘ery sage tthe reproductive el, Grevy' zebra ‘The reproductive organs ofthe Grewy’s aera mane are simular to bat ze about 1% larger than those of phins zebra! Estrus and ovulation Smuts divided the esitous eyele inal zeben species into five phases: proestus, stra, meters, Gliesns, and anestrus? Prowse the phase imme diately preceding estas, and is eharaterie’! by an increase in ovarian activity Mares ding this phase will anevaly reuse stallions attempts lo mount. russ the period of aceptance of the mae, hen a ‘mare wil exhibit the display posture and send for Zebra Reproduction 2855 4 Sallion, A mare inthe display posture stance with her les sighly apart and he ti ited. The pos. {ure quite conspicuous, even fom a distance, an seas fo bean important ateatant fr the salon = Inaeased interest ofthe salon the openmuthet facal expression of the mace (often called awing), land mating are other diguostic Features of ers! During metestras reprdiuctive activity subsides as the destrous phase begins. Desir fs the period of Tveal att between avulsion eyele,and anes Js he period of sexat rest duning the non breeding, season. For the most part ama in nests does not ‘ovulate and te wer is small and anemic” This fs tenor state of the fily belore puberty, 1 zebra mares, ovulation occurs a the ovarian fossa, a depression atthe fe, ventral edge of the ‘ovary? This structure, present in all equid specie, is lined with germinal els destined to develop ito ‘ora Ovulation occurs only at this steutre and never from anyother region on the ovary In plains zebras (andl prebably mosiinin zebras), the ovarian fossa isnot present at puberty, but develops after fewestrous cyels In Grevy’s eb, the oven fos ‘silly developedat puberty tin oer seepects semlesthatof the other zebra species, Overlan activity during the estrous cycle Plains zebra Ovarian activity during the estrous cycle was out lined by Smuts? using, data collected fron 310 fteeranging mazes shot ducing a game copping ‘campaign inthe Krager National Parke, South Aficx Daring proestrus, there a incense in the flicular sexily, with one folle generally larger than the tes. AS in heres, repressing corpora ften om prev ous cycles may be present The estrous phase it Smif proestrus except that the fli thats gor ing to ruptures clearly idntied and the mare now ‘accepsthe stallion In motets, the Gratin fllile thas mptured, a coxpus hemorchagieu forms, ancl the fliculae cavity ile with a yanph-like fluid. As the cals Tning the ruptured folcle undergo capic tivison and become Iatenized, corps lle is formed. Te carpus luteum atin is naximrnsize ‘dir this period afer which it gradually regress. Foliar activity Is Targely dormant, Should the snarefsil to concave, she enters destns or anestrus During desis, a new wave of flies starts ce veloping and the corpas luteum starts Io reps Diestous is genealy followed by the proses phase. The compus luteum is sill present bythe time {ie steceading cyele commence, na in this espe, the pains zebra maze is similar to the doinestic horse Mares that have failed to conceive by the2856 Mare: Other Equids ao. 8 FECAL PROGESTERONE (ngs) —— g & wan + (660) r010¥189 29094 Ra gure 209, ens of fea progisteone a eal encoun ring sce estou in a Gavys tbh end of the breeding season may cease to yee and {0 into nest The evar uf enestous aes ae Inegly inactive, containing some ral flies at some tiny, egress corpora Inte Mountain zebra ‘A detailed stody of Female reproduction inthe Fat: ‘man’s zebra was conducted by Wesln-van Aare al? Ovaries and plasina were collected rn fre ranging zabra. mares shot in the Etoshe National Park, Namibia, Both proesroue and estrous mates have an average of 168 smaller follies (1-101nn) and 1 larger follicle (ether 21-30 or 31-40!mm) present in each ovary. Corpor tea are not present In dist, compors Int ate formed snd several mall follicles (averaging 19 follies per ovary) are sill present olicular activity is simile anestrous ‘mores, with an average of 20 small flies and 0 lange folcles per overy, but corpora later ate not present, ntermediate-sized follicles (at 1-20mm) representa every stage, Grevy'szebea Captive Gievy's zebra mares st the Basle Zoo, Switzerland, wore receptive for 29 days ina 19-30 ay cycle (ef 39 in et 1) Tn the sty by Asn oat females were receptive for 3 to 6 consecutive days, ‘wih an estrous cyte dration of 28 fo 35 days Sin Say Smuts roported a day estrous phase for mazes ‘n Kruger National Pack, South Afi, ‘Asi et ofS monsured estradiol and progesterone {in fal samples from Geevy’s zebra mares dri consecutive estrous cele to compare with courtship. and mating behavioe Figure 3023), Gestation, Plains zebra ‘Sestation length in captive plane zebra atthe Basle ‘co, Switzerland, was reported by Wackernage?” as 361-380 days, with anaverageo! 71 days, in 28 preg. ancies. The PiadelphisZosloyical Galen, USA. ported gestation lengths of 341 andl 335 days ‘These clan are bx acrordance with those obtained from tee-anging animals by Klingel? at 378 and 85 days, and Sinus? af 398 days, At 6-7 montis {ior to birth, a mare's abdomen is enlarged (when Mew posteiovly) and pregnancy i eviden| 2 Sone pregnant horse mares have been reported (2 Sow ests behavion* which inay alg acca in ‘bras, so calculations of gestation length based on ‘sttous betavior may not be rlabe, ‘Mountain zebra Gestation length in mountain zebea has been esti nated at shout 1 yea In the wild, 8 Cape mountain female was estimate to give birth after 359 day? and a Hattmacu’s female was estinated to give bithater 362 days? captive Cape mountain zebra ‘mate gave birt alter 75 days.” These data te fs, ‘at giten the gestation eng of plane and Grey's 2zabrn the igus are rot unlikely Grovy's zebea Inthe wild, gestation i longer in Grevy's zebea, averaging 990 days." Gestation lengths in eptive animals have been shown fo range fom 35810 438 days by Read et al® (8 Louie, USA) and from 390 to 406 days by both Antonius (Germany) and Kenneth! (England). Dstension of the asdomen tends to be snore apparent in Grevy’s thaa in other zebra species" Gestation length canged rom 391 to 406 days for mature captive mares but was 425 days fora young primiparous mare ‘Ovarian activity during gestation Plains zebra King! investigated ovavian activity dusing gestation Jn20 fee ranging pregnant animals, postmortem. He found that until approximately 22 weeks of prep nancy, corpus luteum of sont 2:5 in diineter was prevent tone af the ovaries. Aer 22 week, i Sega regress at flr all hae semaine as 1 03cm wide sip exending approximately 09cm feom the ovarian fossa. Folicis of about 2c in dt meter were present up to 17 weoks of pregnancy, de- ‘creased to Oem at 48 weeks, and showed renewed ‘sett at fl tm, meassring Lem, Sina? found Smlar seals inthe Keuger National Pak ebro During the fest 190 days of pregnancy, he found a ‘umber of intersting phases of ovarian activity. AC ‘er conception, ovarian activity gradually meresed, reaching a peak between days 40 and 10 For the Int terhall of gestation, flies Inge than 12mm sere laygly absent fom the ovals although smaller r= _gresing follicles wore often qute abundant. Active corpora lutea were a red’browa colo, changing to yellow os they represen, According to Sats activity was not equa inthe ‘nwo ovaries. The et ovary appeared to be more ae: tive than the right In cases where mates proce lleorpas tein oniyone ovary toto 13 were courted in the leftard only 73 inthe right (50% let and 0% right) In adit, the let varies of plains debra mazes were lager, OF 340 ovaries (170 pais) smesured, the eft hada greater mass than the right in 120 instances (0.6% Jet > right), This was la con trast fo King’s ells He found ne iffrenc in c= tity betreen the Ho ovarie, with ative corpora Tes occurring in the right ovary in seven animal, sand in the lft ovary in eight animals Zebra Reproduction 2857 Mountain 2b Dring gestation, ovarian activity inthe mountain zara as been reported tobe much like tat of other ‘avy spaces. Joubert conducted a detailed study ‘on feeranging Hlartmann’s zebra females in which he found that large corpora Intea remained in the ‘ovaries even st advanced stages of pregnancy (up oS morths). The average size ofthe corpus luteum, belore eduction was 25cm. The color changes of the corps Iuleum were the sam another 2abra species, with ative ftel bodies having eon ‘olor betre changing to yelloe with advanced se igyession. Ovarian activity wae reported tobe equal Inthe eo ovaries, with follies maturing in ether the ft o: the ght or more olen in both ovaries, Grevysvebra According to King" gestational ovacian activity in frewranging Grevy’s maresissimilar to that of plsins zebra and mountain zebra Uni about 24 eceks of ‘rognaney, 2 corpus luteum of about em in dine- tor wae present inane af the ovaries. By 34 weeks, he corpus eum hed regessed to about tem, bat per Sibel at Io.em upto fll ez, This persistence of 9 fairly large corpus luteum cing pregnancy Was nck serie with endovine atv tl pgs content othe corpus luteum doceased fom 4750 at B weeks t 500g at 24 weoks Follicular sciity wa consberable ducing early pregnancy (Sem), but Ina ceased by 34 weeks Follies at Lem in diame: fee reappeared at 2 weeks ad reached 3c by fll seam. AS wth plains ana mountain zebra setive fl cles wereadaek red-brown coor andl changed to yee low asthe regresied. For Grevy's females, ovarian activity appeared tobe equ in the to ovaries with cogpora tea forming in both the Tet (ee annals) and right (hue animals ovaries "Astet”# measuzed levels of estrada and proges ‘erone excreted in feces uoaghout pregnancy (ge ture 3024 and found patierrs simular to those re ported fo: horss (eg, Squires etal). They alo measured equine chotiote gonadotropin (eC) in trie, which also revealed a pattem sala to that, eon in arses Levels were in the detectable range beginning between 35 and 40 ays gestation and e- tured tg baseline between days 195 and 200. Rar- ‘ny tl also reported eC elevations during that petiod of gestation. Parturition Plains zebra Zebra mans give birth in mach the same way a8 co- reste hoes. Wackernagel> reported infornation2858 Mare: Other Equids FECAL PROGESTERONE (099) _g 8 g (69) o10¥u1sawo88 & + tuo B68) 908 may JUN JUL AUG SEP OCT NEV DEC JAN FEB MAR APR MAY ‘gate S024 Paterno ex prgetone nerd and ary sine chosinepomadtropn concen ing so ‘epi siayellowed by Penny na rey en! Ue corns papi from captive animate, dated account in the wild vas recorded by Klngel!The mare was recur bent, yng Hat on her sie. The hea ond lon legs ‘of the foal had been explled and rvere completely ‘covered bythe amrion. Parturition wns completed 2 mints neat which Une the mae sa up The fal tempted fo stand saveral Kes, breaking the am oti sc and umbilical coed. Aer 14 minutes ad four attempts the foal stood succesflly. The mare Ticked at and chewed the anotic anc but di not feat any of The fal began to wal sbost, paling fot the plicenta In ther instances observed, the penta Was retained for more than 1.5 hours. Aer Parturition, the mare and foal slo roined their Ihaern group, which ignored the bicth and the pres. ence ofthe newborn foal. ll instances recorded, the haem male remained within the viiy af the foaling mare, betiveen 10a SO meters Singlebinths ae the nocmal expectation although at lest one in- stance of aborted twins has becn reneded.® These Accounts are consistent with the these stages of Ie bor recognized in domestic equids,** chaacterie by behavioral eslsseess and the expulsion of the amiotc fd isthe inal stage, more predominant bsominal contractions an the expulsion of the fal {nthe second stage, and finally, expulsion ofthe pax cent in the thie stage Mountain zebra Pattusiion has not been witoessed in mountain = bras but is likely similar that of other zebra species Grevy's zebra ‘As parturition approsches, a Geevy’s frale leaves the herd for the saely of heavier bush Birth nor mally tas place lying down, with the foals hooves Appeecng fs and usually takes only about 7-8 min ‘les, The fal foes elf om the amnion ad eas towards the mothen’s heed. Phe anther licks the fal sean and unlke a plains zebra, ingests the tem branes and some of theamaiote Raid which may ‘be Important in intilzing lactation or the mother Jane bond in thi spec, Postpartum estrus Plains zebra ‘Kingerocorded the shortest interval between to Dints as 378 an 385 days. The latency to foal heat has been repetod! to vary kom 7 #0 10 days after parturition This combined with copulations by ‘mares when their foals ae only days ol, indicatesthat plains zebra exhibit an estrous pedi anya few ays aforfosling Remnants of « corpas Iuteuy, slighty lager than thatseen at fll tern averaging 258m), were fount ln the ovaries of a Hnctmann’s 2ebra femal 3 to 6 wecks postpartum This suggests an estrus 1 week after giving birth. Observations of animals at the Daan Viljoen Game Reserve, Nanbin sovred the foaling-conception interval to vary between 3 and 7 wees, indicating that conception doesnot gener Ally take place during the fst postpetam ests in ‘mountain zebra mares Grevy's zebra According to King! freeranging Grevy's zebra mares showed a range of reproductive atvity post partum Soine animals exhibited evigence of esos ‘eles from 2 to 16 weeks posipatum without con. ception, while others conceived between 3 seeks fo {months postpartum. According to captive sacs, at east one animal hasbeen cored a coming into tstrus about 7 days alter giving birt” and past us has boo estate! to It fon 9 40 ee iaaays Stallion jehavior In all ce species of zebras the veprcctve be havior of ales involves ntoretons with emis as svellas breeding and bachelor mals. To identify aad locate fomales in ests, males will often fehmen to ‘scens depesitad on the grou by the females, Once ‘tmale has four a female be wil atempt to most her if succossfal he will consort withthe female for varying dagroes of ime depending upon species teractions with bachelor mals are typically ages sve, bat depending on species, conperationainong ‘redingstalions is necessary to reduce cuckolding ek” Plains zebra hans zobeas live in muslitiered societies. One level conssis of cove socal groups, rich as closed member ship harem groups and lore associations of bach lors, while another higher level, consists of herds comprising harem and bachelor groups that come. together and break apart relatively frequently! Zebra Reproduction 2859 Hirds fom where resource abundance is high tnd where bachelor associations are mimes ane sing. By banding together, saline amortize the ‘cls f delving away coordinated groups of cuckold + ing bachelors while providing thei females ith ‘crased quality fening time. Meuetain zebra Lite plains zcbras, mountain acres also have closet membership harems and bachelor groups ad herds wil form. Although no details of the social dynamics of mountain zebra hor exit ely that hex dynamics ae similar plains zebras Grevys zebra ‘The open membership nature of Grevy's zebra groups evens that relationships within and between the sexes are transitory. Males establish terstries ‘where ll matings occur Males, however, offen lenve the tertitores when na females ate resent Isle ‘of searching for females, these males are searching {or bachelors By associating withbachelors at vine forcing their dominance of teritry terstrial males swe noel abo aspnce bachelors whe they ‘are presen on estoy with females ™ Also when on temory stallions atta o eesti estrous females from leaving ther teritores, Males snout hem quently and ejcalte at hourly intervals, fh presious sperm from a female's reproductive tract. “This form of spenn competion i aise! bythe fact hat Grevy's zebra maleshaveaisproportonately laoge estes or their body see, Seasonality Plains zebra Aut plains zebra stallions in the Kruger: Na- onal Park, South Africa, show a greater degree of ropredocive activity during the sumer month prteulorly tn January? "This coincides with the ‘mares’ activity pater where 85% of al fal ace por dace ring the sumer months, again witha peak in fawary™ Testicle mass and seminiferous tubule diameter increase about 24% ant 13%, aespectvely, from the months of lowest breeding activity tothe peak months. Spermatogenesis never completely sent, regardless of activity level Desplle test peaks inactivity its cleae that plnins aba ae ess Seastul than other Asean mammals, af foals are produced throughout the eae.2860 Mare: Other Equids Mountain zebra ‘Mountain zebea stallions exhibit increased reproduc tive activity around the rains. At both the Mountain “Zebra Nationa Park (MZNP) ane De Hoop National Reserve (DHNP), South Attica, Cape mountain ‘zcbra stallions show inereased repodctivesctivily daring the spring, with the majorlty of foal being produced the next spring/summer when the anes fe flush with resowres from either the sumer (MZNP) oc winter (OHNR) tains? Similay, Wistlirvan Aasdle et al2? reported that in Etosh [National Park, Namibia, he majority of eproduct activity in Hlarmann’s sea stains coincides with the start ofthe ala, occuring between November and April witha debite penkinFebevary2™ Grewy’s zebra ForGrevy’s zebra in nore Kenya, most repr: "ve activity cccurs early nthe ong tans of ne and July, with another, smaller peak of activity nthe short tins of September and Otter Tis behavior cine es with the onset of estrus in Grey's mes, then they seek out and attract males. Puberty and breeding life Plains zebra Puberty inthe zebra station can be dined a8 “the Interval of time during which reproduction is pos sible, but terminating when fll reproductive capa bily or sexual maturity is reached (but see Skin- net). At the Ngorongoro Crate, Tanzania, Kingel? repoced the youngest haze talons fo be between San 6 yoarsold, with most ania letving thera tal groups between the ages of 12 and 25 months, al ‘hough someremalned considerably longer (up 4045 year of age). In Kruger National Park South Ain the youngest hares stalion was 4 yeas oly and 80.8% ofall harem saline were between 45 snd 12 years ob. According to the Reghonal Stulbook for North American Zoos, the youngest ste wap only 8 months old when the mare conceived and another was only 1055 months of age Cnble 02.3," ‘Accotling to obsavatons of plive anil stal- lions are physiologically mature at 3 years of age. Inthe wild, theatfaineet of puberty seems tobe a _gratual process and somal maturity omnes ter? Par erty was estate tlt frm about 25 tS years of age. Spermatozoa were present in small nubers in animals aged 25-3 yeas, bat appreciable ni bets were not present unl 5 yeas of age: Males ‘were not socaly mature (able to atract an aque females) until 4-45 years of age. Tester mass in essed mot dramatically beeen the ages of 3 ant 5 years old (rom an average of 65g fo 302), while tubule development ws more gral, tasting be. fore the anda frst yen ot an average of 40 diameter, a canting int edulthood (oan se ‘rage of 175 in lamer) Upon eaching sexi maturity atabout 4-45 yen old the combine testes nus reached 80g ane hubule development became selatively conta ‘Sima indicated thot prime breeding silions are tinder 12 years of age, but breeding doesnot stop i ‘older animals, as indicate by King! Salons over 18 years sil ead haeems, The eldest ead stallion recorded by Smuts was 21 years old Theoiest male to ite young in North American 2005 was almost 27 years ll when the mare conceived (Tale 3021). Mountain bra Puberty in Cape mountain zebra typically begins at about 410 45 years old, wth most male attaining females at about 5 yous of age A detailed study of tts as and spermatogees o eanging animals was conducted by Penzhorn and Livy ‘The testes of bachelor males aged IL, 24, and 23 months were prepubelal n stlogtalappeatance ‘tive spermatogenesis in the population began mt about fo 4.3 yoars of age, but this was alleced by Gaylongth. Animals approaching €yens of age dur ingincresing daylength conn showed more > 'ivily than dd animals approaching 4 yeots of age storing decreasing daylongth conditions The social environment was alo important. in the absence of ault males, a Cope mountain zebra colt runnin, ‘with mares sre a foal when only 3 yest old. The tests mass of adult Cape mosatain zebra in MZNP was found to be appredably less (at ~70g) than tht of other zebra species The mean mass of Doth testes plus thelr epididymides war 1654, In ight of ine specimens, the right tests was consi tently heavier than the let This isin concat fo what has been seen in plains 2ebea males? The mean meter of the seminiferous tables was shown to in nse ropidly between 2 and 6 years of age before leveling of In Flarmane’s zebr, the average weight ofthe tess af 87.6 i also markedly smaller than that of the plains or Grevy’s zebra. According to Joubert, tests sie varied with season, eeaching 3 maxim size 094g) in January and a minim sie (8.1) in September. The testes started increasing in Weight i bout 2yenrs of ago reaching die masini 328 stavound 35 years of age.Grevy's zebra Freeranging Grevy's aera males recs puberty at about 4 years af age! At this stage nals ae likely {o be living in bachelor groups, no! consoning with ‘mares and fals until year orto later Captive an. mals in Ragland and in Kenya seichee puberty be- toveen 3 and 5 yeas, but exited! low conception rates unl about 6 years of ge. Reade a however, found that in the USA captive males could achieve sexual potency at agesas Young a7 months, al hat ‘one captive animal beed until 18 yeas of age. More recent data trom the International Stulbook doe rent thatthe youngest male ble tosire young were 2 yeas of ag atte ine the mares cneived, ad ‘hat the oldest was almost 25 years (Table 302) King! stl the tests sion and weight of foe ranging, Grevy’s zebra in some dell The testis of 2 young adult Grevy’s zebra, mentusing about 11585 %75em and weighing about 475g. is sg nificantly larger thon that ofthe pins or muta zebra, Acording Wo King, tests size and weight in the Grevy’s male varied with reproductive ste, A teritorial male's testis welght was recorded a 600, ‘while the testis weight of an older mate associated ‘witha mined herd was rsonded ot S50! The estos terone content ofthese animals algo varied, with the young male. the territorial male. an! the aget male Raving 154 415, ae 595/100 ug of testis respec tively ‘The male reproductive organs lies 2ebea ‘The male reproductive organs of cle enimals were stucied in etal by King! The testes ate carrie the scrotum, sohich hangs between the thighs and posterior to the penis. The deferent duct passes up loom each testis though the inguinal eat to the accessory organs tat le onthe venta for of the pelvic cavity. The ducts (Oem in dininle) enlarge to form the ampulla (12cm in diameter) a6 they ass back over the dorsal surface ofthe Blades al ‘pen atthe beginning of the urethet. These of the seminal vesicle opens into the arethra behind each ampulla atero: to the prostate gland. The prostate sland is lobulated, bilcbed gland extending 25cm ‘oneach seo the wethra. The blbowrethal lane are locate about Sem posterior ant Inter to the terete at the base ofthe penis. The urethra passes round the posterior border ofthe pubic sympysis imo the pens and extends along the venta surtace fof the abdomen, The extremity of the vrethra Is enelosed within the prepuce and lie in the region of the ambitions Zebra Reproduction 2861 ‘Mountain zebra ‘hearst marked entre of the male reprostctive ox- _gansin mount zebra isthe size of the testis, A am laverageo! 70 and 86g for Cape mountain and Hart mannis zara, respectively, the mountain 2ebra testis Jsqppecably smaller than that of ether the pins ot Grewys zebra Grevy’s zebra 'Bven when controlling for animal size, the male 2c- cessor organsare much larger in Geevy's zebra than In he plains or mouatain zebra"? The most con spleucs feature isthe large senina vesicle, which fan contain upto 400 nL of ii in he largest tal lion sampled and upto liter ins captive animal at out 5 years of age. Tests size and weight areal sSlgniiantlygreaterand vary with reproductive state ‘ofthe male (ee above, Foal Plaingzebra ‘Single srs wore corde by Klingel and Kings Inthe wild andoy Wackermagle in eapivity. Ringel caleula ese ats forthe foals born in the {gor Cater, Tanzania. OF the 170 fosls born, 8 vere females andl 74 were males, giving a ratio not a ferent fon 1, Foals extricate themselves from the Dirth membranes unaided, stood within 1 fo 3 aie toy, stpped afer 19 minutes, walked ater 32 min- utes, ar cantor after 4 t9 60 mints. fn the one Instance of actual fang reconted by Klingel> the offering suckle forthe ist ne within 1 how and "more nes tm the next hous for secon 25minutes ate for toll of Miminutes, Wacker raglo® reported rst sucling at 71 minutes alter pa Durtg the fast fw days to wecks ofthe fot the moter is very protective, This coreates wi imprintng peri in which the foal learns fo recog- nize its mother by her call, her scent, anther see pattern. Until impriating is complet, the fd ill follow any large abject including vehicles and the rare’ protectivnessInsuees tht the fo will not Finprinton another animal. Ding this period, other ‘members of the group (lnluding the harem stallion) ste not lowod to approach the fal to closely. Sch behavios insures protection ofthe fs! during spar ticularly vulnerable stage Beyond this period the ‘mother sn foal remain sos, but oes at allowed to interact and socialize with the fos. Mothers with young feasaremore wary than ober members ofthe froup aul are the ist to move ola signs of danger.2862 Mare: Other Equids Mountain zebra ‘Actual bits of movntainzsbrs fos have not been ‘teconled in the wi" but data reganding sexe tio at birth, mara! behaviog, and foal activity do fest. The sex ratio at birth snot different Sram ‘neither Cape mountain of Hartmann’ zebra, an ‘the young ate just as precocious, getting up, walking, and running within a short perio of tne? ‘Dring the first few weeks of lf, mountain se ‘bra foals and their mothers behave very mich Ike other zebra species. Hoth Cape mountain and Hac ‘mann’ zebra foals remain close to thee mothers in thesiest weeks, nthe ir! ew hours ater bith, Hat: man’s zabra mares spend much tine sling and licking the foo. This maybe important nse te female’ recognition of young Both Cape mou fain and Hartmann’ 2bea fons make frequent ta te contact with the mothers, rubbing thei eas along thee mother’ fans, initiating naso-nasa cov tact andl rubling at thee necks In both subspecies the mothers actively prevent contact between the fat tail other members ofthe group. Thiscanbe taken ‘extremes in Hartmanw’s zebras sone mothers will emain separate fom thee groupe for several ayn” As in plans ad Grevy’s ar, this behavior ls cor- related with the ctcl peiod of imprinting of te ‘mare's sound, scent, ana stipe pater on the fod [Mores of both species are mose skits than other group members and are the fist fo flee at any sign of danger Greuy's zebra Grevy’s zebra fouls are similar to both plairs and mountain zebra fons, Single births have been recorded both in the wild! and in captivity? Dat {om captive animals show thatthe young are pre cncous, standing 6-Idminates after bch, walking by 20mintes, and running by ABanisuten At abou 29 minutes oe foal atemps onus ancl does sob (minutes At 97 minutes the fol stands well and at outs nurses wall. In captivity, the sex ati at birth js os? ‘The Grevy’s newbom wil follow any lage, nov ing object. The mother 6 quite protective (particu Iasi during the frst 2 days) an keeps other aniale way until er offspring has printed upon her cal ‘con and stripe pater. After imprinting hs been achieved, the moter and foal wil joins group of rursig mothers. When traveling to water, the mare will sometimes eave the foal with his group. Weanling Plains zebra ‘According to Smuts? lation generally continues ttl T te 2 months before the bitin of the nent foal. ‘The eariast weaning occurred at 11 months of age, butif amare does nt eoneaive soon afer faling, she will continue to suckle her foal tronghot and be- yond is first year One captive are tu itwas tS mandhe ol, weaning fothebirth of her nest fort, ‘Mountain zebra (On average, meuntaln sera foals ace weaned at shout 10 months of age shout 35 to 6 months be- fore the neve foal is born. For Cape mountain 2e- bra the ming of weaning ie highly dependent on the bitth of new offspring and can be gute vacable, but for Hartmann’s zebra, 10 months seems to be the morn. If maves do not conceive duting postpar ua ests, sickling may be permite fr longer pes ‘ods. Penzhorn®*? reported infrequent stkling in animals oll a 2 months oF age Grevyszebra Grevy’s zebra foals are wenned at about 9 months (62.42 weoks), but will remain with their mothers for several more months or yeas, depending on the sex (se below). The young ae sem independent at out 7 montns, often being let by the mother ot leaving her fr hous at 3, Yearling Plains zebra Young mares a the age of 1-2 years, aroun the time of ther fist heat re abducted fom thee fares by stalions.Klage™repoted tht on average, mares were abducted at about 18 month but as oun a8 19 months of age These young fertales do not typi. cally remain withthe ial stallion, but within aout {months of separating from their nat group, Ie come permanent members of eproduetive group, ‘Most young salons leave the natal group, at aout 12-25 months of age in some cases ater, ad either stay single or jin other stallions to form bache- lor groups, Bachelor groups ae of vacable umes, up fo 18 animals. smaller groups of two, three, anc four have been reported tobe very stable. Departure ‘ofthe young males isnot cvtlat with sexual ma- tury or any antagonism with the lad stallion Put seume tobe detormined more by the composition of the natal group. In parila the males tee To leave ster the birth oftheir mothers net fal Mountain zebra ‘The timing of dlispesnt is similar for mates an females for both Cape mounts and Hastaann's2sbras. In Mountain Zebra Nations Pare (MZNE), sale ane female Cape mountain rabra left tele ratal groups at an average of 2.85 months 52? in De Hoop Nature Reserve (OHNE), however, females leave their groups calor at 18.85 months, than do the mals, a 24:36 months” Female dispersal coin. ices with estrus onset, but there is no physiol ‘correlate in mates. bot MZNP ant DHINR. the seating eave the group oftheirown acord and are not freed todo so by either the stalin o the mare Tadd, dupereal snot conlted with he moth cer subseguentpregaances othe stats.” and sal Tionsin MZNP wil vigoonsly attept to retain thee yeasingofopring.®” Dispersal in young movin 2a is ikly due to bond taneevencs fom mater ral/pateral to age cohort sesociations, Ts i Tae to what Kling report in pains ze Instark contest, Hartman’ zebra typically leave ‘he natal group at around 14 to I mons age” At ‘his stage Shorly before the bcth of hee ext fol), the mare actively testo drive the older offpring (ovhether male or fem) oxt of he group. Appran ‘mately onesie of yearlings leave the group a8 ade ect rsul ofthis aggressive behavior The emailing tworthieds leave of thee oven acon, sometimes 38 ‘uch a years ater. Upon de birth of her next fa, the maze will ignre the Yearling fit has nat alteady Young Cape anountain mazes often jain boche- tor groups before joining ote stable, reproduc: tive group. This was alays the case for mares i HIN! anv vory often the case for mares in MIZE At DEINE, the females stay in the boc lor groups approximately 87 months, leaving when they are about 289 months ol. The ferales are not abducted by harem stains they leave of thee of accord. AU MZNP, females wil sometines remain ein for severa ont befor ning either bach ‘lor oF reproductive groups, Okler mates ae taken ‘up by harem salons sooner, while younger mares re either salary or in bachelor groups for Longe petiods. ‘Young Cepe mountain males in DHNR stay ta their natal groups longer then the ems, fr ane ezage of 2435 months, leaving when they ae about 52:4 mouths of age" These data agree ith Pen 2hoen’s estimate of the age at sential mati. Ia "he MZNF, young inales roam singly or in neable groups only local associated with the more sable Dacheloe groups” Affiliation withthe nat group i song, young males having left after the displace nent oftheir fathers wil often join them. In ation, young males having let hel natal group sil ntact ‘will often rin the group fr short periods artmane’s zebra females are typically herded by either bachelors, forming the coe anit of new reproductive group, of by harem stains, therby Zebra Reproduction 2863, Integrating into an. aeady stable reproductive |roup. Young males leave thet natal group (0 join Dacolorgeoups that eorsst of mosthy young, bat alsc older csplaced talons” Grevy’szebea Yourg mares lave their mothers at about 13-18 smaaths of age, typielly 2-7 months into the io zs nox pregnency Te females wil not breed n= til they are between 2 and years old, Dung their fests a sour years of yp) te mae te, attract sepatate them fom their mothers to mate ‘Move experienced mares till pretent only to tet ‘oval males, cousin Tess disruption tothe female scape Crovy’s mals slay with their mothers wnt they reabout’3 years ol after which hey jin a bachelot group oramied her? Young males donot ypially [prinaces to emuales nll they are about yensold, sp ave abachelor period of 2105 years® References 1, King In le ide oe repo fhe Cans sepa Cy nen at te ra © Sahat) fone Kaper tonal Pah ae fievraioanse ™ Saute GL epedctin tthe 30et stiin (Eg Dt agsaran om he Eger Nar Pa Zo igs apcomoataan 2 4 Aa CS uma J Heston 9, Pach MT, Ro Bowe en Keer Patras cn a Ciesradet and progeerane se wi Chon ‘eeopn in resets Eqns gg aly Sisand ppc 20 Oey HOES ‘rota Hava A Mao NovenyV. An satan hepolton roo’saa aw gey0in Era Klemm 20 Yorke 8 3 6 Kings 1. Repredactin in the ple sb, Bins Sache boc ene egal cre] pd onl sip ess ashen BL: Repotine haiti lates gg pelt of Cpe nonin bn es ets se red ea e817 ed Kay C Bae Mug Gory tex maa: onsen Ames Ashen of Calg Pk ‘Se Ager Spies Sree Pah 5 ts MO St isi Zea P98 pp. Graber J Rutentin DL Sper cpio ovat ‘inate ng ki Bl el Sc 198, 10. Reena, Hack M,N an sesso an Phe evtaton ome sot nih sae ‘veh comparson to pst, be apps ne wed) Sel St in Pre Ne an Compre espn Catrige Une Pesy, MO pp B=2864 Mare: Other Equids 1. Angel A compton ofthe wil organ ef ‘quta i Proceng of Confront egy od Sorat sas anonyme 1. Robern DE elegy and soy in Hse ad 2 tee. Ratersen 1, Wann 0" (tt gd ‘eS sti Fa Sb of edn evcion, N Picton Unversity Ps 186 po. 22 eo 18, Ritesh I. The ety of fn cl bso in bn onde ot Rose Ae) A al Sot tant, cn oO, ter Kyoto Uaterty Pe pp. 1 14, Rate Dy Ne C. Soci snd podcive se leas pl tran Hg Jos Ct) ope Sen vert Pra nd tonal Ct, Com Iwidge very Pre 2009p. 96-36 138 Gaby JR Te eelgy of eal ts so mae ‘mang Stes inthe Gy srs Sp Za! $e Lone ‘vata. pageant 16, Rowen M Nether fant cir and egy of rey? a Eqs ery New Haan CT Ya Urey PE io 18 10. Lipp PH Rasa OA Sty repratuc suc ond erga note fet) 1. Bch My Sunday 5 Cary J Lakin HM, Ecler My Ruben DL Sclipo: ‘ste eat nce dep mie inovermets pinne he (Equus SP P35 19. Moet J, Pia Zt Noh Aram Rei Sink 20, Panther BL, Lloyd Pt Compson of poine ps ter oo Cope mount sre eg br 7) ppinon | eps 7 13. 21. Joubert E Ne the epodton tm 2 ‘eatpnaebes arma sann Wet Aes Mat brass 12 Wostn-y Aare 1M, an ade BSD. Ree ‘loon a aml tna a, Ee a ‘oJ oot Fre 8865051, 28, Kingdon | Est Ant Monk A of Eoin ‘Aft Chega ie Unto of Cao Pr 1, 2, lan SD: Gays ata ely in eget nvm PRD ten Urivety” Cage Caan ie 295 Langone Ze Inert Sto. Wich ‘site Ue Mel 20208 26. King Nes on te Bigy oft pas sabea Tuts gga token matte Ev Ae WM fo maiseaae 27 Jute The sl orareationand sence eho {nee tigcmaen pn guar een haces ge Boagase 28 Pesen a rMeve Nein a nf bln Cape mount rage ah eed Fen near 29, Boneh, ght) ery Cer. ono: lis “inal nd Cy 22 1. Am C3 Bgl prt isto. Mehinan PD) IcwSSC Ep Aton Pon hind Sern UCN, ‘aurepp. IS 38, Kopler The sent yee domes mam ‘ae Depurant of Apne a ih eps ithe Dito of Yenry Een at Rae 138, pez. 232 Hammond J, Woah K Anam nd Bogie ‘hang ng tsetse eae P ay So Ioulbnw isu {8. Waenapel 1 Goat stn gas bucel och at ase Zeo"scntutanobredng ogy. nero! Zoe sa a Brown CE Reng wil anal captivity and gesaton pds Manel 9361710183 38 Sts CL. Pe and postal growth phenomena of eer buh tio. 7 4 ADNCS Goleta, Gather} Asem he sv Sehr pregnet es Hom Ble sea 1 7. Peahorn BL Saviour and popultn eogy a he Speen ne ni 38 Groves er, Ae te i, Neon Ae et and Loon vi and hl, 9 9, Autos. nie bemecgeato Zac in Shon ‘rs me 1. Zep Cann 305087 10, Kent. Gaon ene Soap UK: Cooney ‘Asestrl ron 95 1. Spies Hl, Doug BH, Selentogen WE Gnker Of ‘Onisian anges dat the estou eye ant pega Innate [nim Sa WALES 1 Rose tote ME Ua Cpt the ae ig peegrseyThrigeney 99H 2 <8 Raat Mar fe ey Byrd ee Rag gsm oni For ay 4. Ginet O} psc Bg te Me ako pe taps dn Cres Pe Wi er 9 16. Tere longed orgnetin and Savin af Cape sain sees Egos bn Pa 2etn fa ppg CesT As 4 Chie Cat ge Momo Sper (7, Kang, Sort RY, Mtn DE Hameron The ep ‘Satine pyoloy a ale abn nena yp Zel Sa on 960931 22 1. Neen ae Fowler ME 6) 2a an WE Aa Mate Panda, OA“ WB Sener aC 86, ppm, 49, Ces I Sec nparznion ad mating sali of Tat tped equ The revys brn, Pera, Ni inten Unies PMD thes 8 A, Bomens CW Psy of gorda hoes id gate cmpoans. Capps CAP) potion Bs rs Landers Acne e190 {1 Shige JD The cnet of pubuty. 5 AP Ma) 1598 52. Trlr: Heotuhngenan den Babine des Gage Toop ep Reo Son 15. Kgl Hy Kings Die Get snes Zebeas (gn nt bet Mate). Zt fr Hepsi iene SL Hh Sundaresan Sh atin HM, Contig JE, Sor My brat DL Aen baer eral pn bra gas bots Eh 2028201 5,55, Jha ES an graces pean pos ‘Sevelopeent of th Hart cn Eqs aie ‘ne Me 78 8, ‘54 er RF thay of ln New York Fou Pes rd Zebra Reproduction 2865 7. Behar BL Oberon of moti f ee saying Cap eatin ge ta 8 0 | Greer CS. Gino's the er 26. St 198512