MACRORHYNCHA AND MABUYA AG/LIS) IN A BRAZILIAN RESTINGA HABITAT

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1 HRPTOLOGCAL JOURNAL, Vol. 9, pp (1999) RPRODUCTV TRATS OF TWO SYMPATRC VVPAROUS SKNKS (MABUYA MACRORHYNCHA AND MABUYA AG/LS) N A BRAZLAN RSTNGA HABTAT CARLOS FRDRCO DUART ROCHA AND DAVOR VRCBRADC Stor d cologia, nstituto d Biologia, Univrsidad do stado do Rio d Janiro, Brazil Th rproductiv cycls, fat body cycls and som lif-history traits of th sympatric viviparous skinks Mabuya macrorhyncha and M. agilis r compard in a sasonal "rstinga" habitat of south-astrn Brazil. Both mal and fmal rproductiv and fat body cycls ar vry similar btn spcis, ith gstation lasting 9-12 months and parturition occurring during th arly t sason. Clutch siz of M. macrorhyncha as smallr than that of M. agilis. Fmals matur at a largr siz in M. macrorhyncha than in M. agilis, but mals of both spcis appar to matur at similar sizs. n both spcis, fmals ar largr than mals, but th lattr hav proportionatly largr hads. Rproductiv traits of M. agilis ar typical ofnotropical Mabuya, but thos of M. macrorhyncha hav som pculiaritis, on of hich (small clutch siz) is blivd to rsult from constraints imposd by its morphological adaptation (i.. rlativly flattnd body plan) to bromlicolous habits. Ky ords: Rproduction, lif history, Mabuya, Brazilian skink NTRODUCTON Th gnus Mabuya is on of th most spcios and idly distributd gnra of th Scincida, ith about 9 spcis found ovr most of tropical Africa, Asia, and Amrica (Shin, 1985; Nussbaum & Rax orthy, 1994). Not surprisingly, this gnus is also xtrmly divrs cologically, ith spcis utilizing a id varity of habitats and microhabitats (.g. Huy & Pianka, 1977; Castanzo & Baur, 1993; Nussbaum & Raxorthy, 1994, 1995; Avila-Pirs, 1995; Vrcibradic & Rocha, 1996). Concomitant ith such gographical and cological radiation, lizards ithin this gnus hav dvlopd a considrabl divrsity of rproductiv charactristics in diffrnt rproductiv mods that includs both oviparity and viviparity (Fitch, 197, 1985; Vitt & Blackbum, 1983; Shin 1985; Blackbum & Vitt, 1992). Although both viviparous and oviparous Mabuya spcis occur in th Old World, only viviparous forms ar found in th N World (Fitch, 1985; Shin, 1985; Vitt & Blackbum, 1983, 1991; Blackbum & Vitt, 1992). Actually, N World Mabuya sm to constitut a monophyltic group drivd from an African linag (Grr, 197; Shin, 1985; Baur, 1993) and, apparntly, all of its spcis possss a pculiar typ ofviviparity charactrizd by an xtrm dgr of placntal nutrint transfr that is uniqu among squamats (Shin, 1985; Blackbum & Vitt, 1992). Th grat divrsity of rproductiv faturs among Mabuya spcis maks th study of rproduction in lizards of this gnus vry informativ, as many qustions about cological and volutionary rproductiv rsponss may b addrssd. Spcial intrst should b paid to th analysis of rproductiv pattrns of sympatric Mabuya, from Corrspondnc: C. F. D. Rocha, Stor d cologia, nstituto d Biologia, Univrsidad do stado do Rio d Janiro Rua Sao Francisco Xavir, Maracana , Rlo d Janiro, Brazil. mail: cfdrocha@urj.br hich prsnt diffrncs may highlight thos historical forcs that may slct for crtain rproductiv charactristics. ndd, diffrncs in rproductiv charactristics among sympatric and allopatric populations of congnrs hav bn rportd for this gnus in Africa (Huy and Pianka, 1977; Pianka, 1986). At many localitis along th coastal sand plains of south-astrn Brazil, hich ar charactrizd by sanddun habitats (rstinga), to spcis of Mabuya usually occur sympatrically: Mabuya macrorhyncha and M. agilis (Araujo, 1991, 1994; Rocha & Vrcibradic 1996 Vrcibradic & Rocha, 1996). Som information ' on r productiv traits xists for th formr spcis (Vanzolini & Rbour;:as-Spikr, 1976; Rbour;:as Spikr & Vanzolini, 1978; Zanotti, Sant ' Anna & Latuf, 1997) and virtually non for th lattr. Although living sympatrically in ths habitats and bing similar in body siz, ths spcis diffr markdly in morphology and microhabitat us, ith M. macrorhyncha shoing som tndncy to scansoriality and living mainly on and among ground bromliads. On th othr hand, M. agilis is a ground-dllr hich basks and forags on laf littr (Rocha & Vrcibradic 1996 Vrcibradic & Rocha, 1996). Th comparativiy mor flattnd body and had, and longr fingrs in M. macrorhyncha hav bn suggstd to b adaptativ traits rlatd to its habit of living among bromliads (Vrcibradic & Rocha, 1996). Flattning of th body plan to suit cological spcializations (such as lif in rock crvics) may impos som rproductiv constraints to fmals of som lizard spcis (Broadly, 1974; Vitt, 1981). W analysd th rproductiv cology and sxual dimorphism of Mabuya macrorhyncha and M. agilis, spcifically addrssing th folloing qustions: (l) Do th to sympatric spcis diffr in thir rproductiv and fat body cycls at Barra d Marica? (2) Dos littr siz diffr btn spcis and is it corrlatd ith f-

2 44 C. F. D. ROCHA AND D. VRCBRA DC mal body siz ithin spcis? (3) At hat siz do mals and fmals attain sxual maturity in ach spcis? ( 4) Within spcis, is monthly variation in fat body mass mor conspicuous in fmals than in mals, considring th lo nrgtic dmands of sprm production (s Blm, 1976) compard to gstation? (5) Ar th spcis sxually dimorphic? MATRALS AND MTHODS STUDY ARA AND CLMAT Fild ork as carrid out at th Barra d Marica rstinga (22 57' S, 43 5' W), 38 km ast of Rio d Janiro city in th Rio d Janiro Stat, S Brazil. Rstingas ar Quatrnary sand-dun habitats covrd ith hrbacous and shrubby vgtation, common along th Brazilian coast (Suguio & Tsslr, 1984; itn, 1992). Th ara has markd tropical sasonality (Fig. 1 ), ith a t sason btn Octobr and March and a dry sason btn April and Sptmbr (Franco t al., 1984; Rocha, 1992). Th man annual tmpratur varis btn 22 and 24 C and th man annual rainfall rangs from 1 to 135 mm (Nimr, 1979). COLLCTNG MTHODS AND ANALYSS Lizards r collctd monthly from May 1989 to April 1992 ith an air rifl. Shots r alays dirctd to th had and nck of th lizards, in ordr to kill thm immdiatly. ach lizard as thn immdiatly transfrrd to a plastic sac containing cotton soakd in thr. This as don in ordr to quickly anasthtiz lizards that may not hav bn killd instantanously and to nsur a painlss dath. Shooting as by far th most fficint mthod found to collct Mabuya in a rstinga ara charactrizd by larg patchs of vgtation. Catching thm by hand or ith lastic bands hn thy ar basking is vry difficult (M macrorhyncha usually basks on th spiny-dgd lavs of ground 14 llo...j...j o ci: ll z ci: iso a:: z J M A M J J A S O N D u a:: :: l i: a:: Q. 3 ::;;; - 2 FG.. Avrag monthly long-trm raifall and man tmpratur of Barra d Marica, Rio d Janiro, Brazil. xtractd from Rocha ( 1992). 1 bromliads) and noosing thm is practically impossibl (th lizards ar vry skittish and usually rtrat to th intrior of thickts of vgtation or bromliad patchs hn thy sns any disturbanc). Th charactristics of th study ara also mak th us of pitfall and drift fnc traps not fasibl, sinc th patchs of vgtation ar too dns compard to anothr rstinga aras hr hav succssfully usd such tchniqu to catch skinks (s Vrcibradic & Rocha, 1995 for mor dtails). To nsur sufficint data r collctd for statistical analyss, th numbr of skinks collctd pr month usually rangd from on to tn (Sptmbr 1991 as an xcption). W bliv th impact causd on th skink population as ngligibl, sinc sampld only a small portion of th availabl habitat during th study. Also, on subsqunt visits to th ara, aftr th monthly collctions had casd, did not not any visibl dcras in th frquncy of skinks sightd pr day (both spcis ar fairly abundant in th ara). ach collctd lizard as ighd (to th narst.1 g) ith a Psola balanc, prior to fixation in 1 % formalin. Th snout-vnt lngth (SVL), had lngth (HL), had idth (HW), mouth lngth (ML) and had hight (HH) of ach lizard as masurd using vrnir caliprs (to th narst.1 mm). Spcimns r thn dissctd for sx dtrmination and xcision of rproductiv organs (including mbryos) and fat bodis (th f lizards hos organs r damagd by th shot r not considrd). W countd and masurd ovarian follicls, oviductal ova and mbryos of ach fmal, for both spcis. Th rproductiv stat of ach fmal as assssd according to th folloing catgoris (modifid from Pattrson, 199). Stag : no yolking follicls; no ova stablishd in oviducts; Stag 2: yolking follicls; no ova stablishd in oviducts; Stag 3: ova or mbryo sacs (lss than 4 mm in diamtr) in oviducts; Stag 4: mbryo sacs mor than 4 mm, chorioallantois stablishd, mbryos undvlopd; Stag 5: mbryos occupying 2: 5 % of mbryo sac; ys and limb buds (or limbs) vidnt (Stags 3-36 ofdufaur & Hubrt, 1961 ); Stag 6: ll formd (nar-trm or trm) fotuss (Stags 37-4 of Dufaur & Hubrt, 1961) Fmals r considrd rproductivly activ if thy containd ova or mbryos in th oviducts (stags 3 to 6). Man brood siz as stimatd for ach spcis using data from all fmals containing oviductal ova or mbryos. To valuat th xtnt to hich fmal body siz affcts brood siz, prformd a linar rgrssion of brood siz on fmal SVL. For ach mal, rcordd th longst and shortst axs of ach tstis and stimatd tstis volum using th formula for an llipsoid (Mayh, 1963). To assss rproductiv condition of mals, paraffin sctions r takn from th middl of th lft tstis (including th pidydims) and staind ith hamatoxylin and osin. Mals r considrd rproductivly activ if sprmatozoa r prsnt ithr in tsts or in th pididyms.

3 RPRODUCTON OF SYMPATRC BRAZLAN SKNKS 45 All linar masurmnts of gonads and mbryos r takn ith digital caliprs (to th narst. mm). Th combind mass of both fat bodis (for ach sx, in both spcis) as rcordd using a Mttlr ltronic balanc (to th narst. g). To assss th monthly variation in mal tstis volum (xprssd as th avragd volum of th to tsts in ach mal) and in fat body mass for both sxs, calculatd th rsiduals of th rgrssions of tstis volum on SVL (both log-transformd) and of fat body mass on SVL (idm), rspctivly, and took th man valu (plus 1 SD) of th rsiduals for ach month (only sxually matur lizards r includd for this purpos). Rsiduals of fat body mass of adults r corrlatd ith rsiduals of man tstis volum (in mals) and ith man mbryo sac diamtr (in fmals) using rgrssion analysis, to valuat th dgr of usag of fat rsrvs throughout th rproductiv cycl. For M. macrorhyncha (th monthly sampl sizs of M. agi/is ar too small), th ffct of thr nvironmntal variabls (total monthly rainfall, man monthly tmpratur and photopriod) on man monthly tstis volum (xprssd as man valu of rsiduals; s abov) as analysd by rgrssion analyss. Du to th lack of publishd data on th tmporal gap btn nvironmntal changs and changs in lizard tsticular activity, assumd a tim-lag of to months bcaus such physiological rsponss to nvironmntal variation ar unlikly to b immdiat. Data on avrag monthly rainfall and man monthly tmpraturs for a 3 8-yar priod ( ) r obtaind from th Dpartamnto Nacional d Mtorologia station of Nitr6i, locatd a. 19 km st of th study ara. W tstd for intrsxual diffrncs in lizard SVL ithin ach spcis using on-ay analysis of varianc (ANOV A). HL, ML, HW and HH r compard btn sxs in ach spcis through analysis of covarianc (ANCOV A), using SVL as th covariat. To incras our sampl sizs for analyss involving clutch sizs and morphomtric variabls (including SVL), for both spcis, includd data from lizards collctd sporadically in Barra d Marica both bfor (in 1986) and aftr (in 1995 and 1996) th study priod. Dscriptiv statistics ar xprssd throughout th txt as man ± standard dviation. Nomnclatur of othr Mabuya spcis mntiond in this papr follo Avila-Pirs (1995). RSULTS FMAL RPRODUCTV CYCL Brood siz of M. macrorhyncha avragd 2.66 ±.63 (rang 2-4; n = 38; Fig. 2a) and as significantly corrlatd ith fmal SVL (r2 =.258; Fl,35 = ; P =.1). Th smallst fmal M. macrorhyncha containing oviductal ova masurd 59.9 mm in SVL. Anothr fmal of th sam siz (collctd on 25 Novmbr 1996) had undvlopd oviducts and containd no vitllognic follicls in its ovaris, hich suggsts that B 1 8 ('.) z 6 :: M. ag ilis 3 4 BROOD SZ 5 6 FG. 2. Frquncy of brood sizs (xprssd as numbr of broods for ach b rood siz) among M. macrorhyncha (a) and M. agilis (b) at Barra d Marica, Rio d Janiro, Brazil. it as not sxually matur. Thr fmals containd mbryos/ova in diffrnt dvlopmnt stags simultanously (rspctivly, in stags 3 and 4, 3 and 5, and 3, 4 and 5). To fmals (SVLs = 68. and 72.8 mm) containing ll-formd fotuss r collctd in lat Octobr 1991 (Fig. 3a); anothr fmal collctd on 5 Dcmbr 1996 containd thr ll-formd fotuss. Th smallst individual in our sampl, a mal (36.3 mm SVL; umbilical scar prsnt), as collctd on 13 Dcmbr Yolking ovarian follicls of M. macrorhyncha rangd in diamtr from.9 mm to 2.2 mm; th smallst oviductal ova r 2.3 mm in diamtr. Thr ll-formd fotuss from a fmal collctd in Octobr 1991 rangd from 24.9 to 25.7 mm in SVL. Ovulation apparntly occurs from Dcmbr to March and implantd ova undrgo littl incras in siz until about Jun, hn mbryos bgin thir rapid dvlopmnt phas until thy ar rady to b born, about Octobr-Novmbr (Fig. 4a). Brood siz of M. agilis avragd 3.5 ± 1.4 (rang 2-6; n = 8; Fig. 2b) and as not significantly corrlatd ith fmal SVL (r2 =.131; F, 16 = 2.4 ; P =,.14). Th smallst fmal M. agi/is ith oviductal ova had a SVL of 49.2 mm. Fmals ith ll-formd mbryos r collctd in mid-sptmbr 1989, arly and lat

4 46 C. F. D. ROCHA AND D. VRCBRADC A (/)...J 1% 6% (.!) g 4% a? 2% a. Mabuya macrorhyncha !l!JJ Stag 1 ZJStag 2 mstag 3 DStag 4 Stag 5 Stag 6 al 2 15 D: (il m G : llj 1 L D: llj 5 - llj : < s % J F M A M J J A S N D b) 8 1% (/)...J u. u. 6% (.!) g 4% u a: a. 2% Mabuya agilis !l!JJ Stag 1 ZJStag 2 mstag 3 DStag 4 stag s Stag 6 15 D: m : llj 1 u. D: llj 5 - : < G c J F M A M J J A S O N D FG. 4. Monthly distributions of mbryo siz (xprssd as maximum diamtr of th largst oviductal ova or mbryo sac for ach fmal) for M. macrorhyncha (a) and M. agilis (b) at Barra d Marica, Rio d Janiro, % J F M A M J J A S N D FG. 3. Prcntags of sxually matur fmal M. macrorhyncha (a) and M. agilis (b) in ach rproductiv stag (s txt) for ach month during th study priod (May 1989-April 1992) at Barra d Marica, Rio d Janiro, Brazil. Monthly sampl sizs ar xprssd by numbrs abov bars and rprsnt poold data from diffrnt yars. Sptmbr 1991, and lat Octobr 1991 (Tabl 2), and r all largr than 7 mm in SVL. Th thr fmals in rproductiv stag 5, from Sptmbr 1991 (s Fig. 3b) rangd from 63 to 68 mm in SVL.Th smallst individual in th sampl, a fmal (SVL = 47.6 mm) as collctd on 13 May 1991 and vry young individuals (SVL :::; 45 mm) r sn in th fild on 3 Dcmbr On fmal containing nlargd follicls (about to b ovulatd) and no implantd ova as collctd on 28 Novmbr Volking follicls rangd from 1.1 to 2.2 mm in diamtr and th smallst oviductal ova masurd 1.5 mm. Four ll-formd ftuss from a fmal collctd in Octobr 1991 rangd in SVL from 24.7 to 26.7 mm. Ovulation in M agilis appars to bgin in Novmbr and ova apparntly stay rlativly small until May-Jun, hn rapid mbryonic groth bgins (Figs. 3b and 4b). mbryos ar ll dvlopd by Sptmbr-Octobr and parturition apparntly occurs mainly during Octobr-Novmbr. Brood siz of M agilis as significantly gratr than that of M macrorhyncha (ANOVA: F1 54 = 14.15; P <.1). Most (a. 9 %) of th fma ' s containing oviductal ova or mbryos, in both spcis, also had vitllognic follicls in thir ovaris. TABL 1. Valus of morphological charactrs of Mabuya macrorhyncha at Barra d Marica, Rio d Janiro, Brazil. F-ratios tst for diffrncs btn th sxs using ANCOV A, ith SVL as th covariat. * * * P<.. Had Width Mouth Lngth Had Hight Had Lngth mals fmals mals fmals mals fmals mals fmals 55 n Man SD Rang F-ratio 28.71*** 63.36*** 17.77*** 78.87** *

5 RPRODUCTON OF SYMPATRC BRAZLAN SKNKS 47 TABL 2. Valus of morphological charactrs of Mabuya agilis at Barra d Marica, Rio d Janiro, Brazil. F-ratios tst for diffrncs btn th sxs using ANCOVA, ith SVL as th covariat. ***P<.1. Had Width Mouth Lngth mals fmals mals fmals n Man SD Rang F-ratio 35.72*** 25.48*** MAL RPRODUCTV CYCLS Th smallst mal M. macrorhyncha contammg sprmatozoa in th lumina ofsminifrous tubuls and/ or pididyms (from 28 Octobr 1991) had a SVL of SS.9 mm. Hovr, both this individual, and a slightly largr on (S8.4 mm SVL; collctd 29 Sptmbr 1991) containd vry f matur sprmatozoa. Th monthly distribution of rsiduals of th tstis-volum - SVL rgrssions (Fig. Sa) shod that tsts ar at thir largst from Novmbr through January, dcras in siz from Fbruary onards, rmaining small until August or Sptmbr, hn thy bgin to nlarg again. Th smallst mal M. agilis (collctd on August 1986) masurd SS.S mm in SVL and had its tsts and pididyms filld ith sprmatozoa, indicating that it as sxually matur. Th monthly tsticular cycl in al 3.. :: 2 iii 8 :: 8 :: :: 8-1 ) - -2 ) - -3 b) 3 "' ::!: ) 2 a: G :: :: -1 ) ) - -3 J F M A M J J A s N D FG. 5. Monthly distributions of tstis volum (xprssd as th rsiduals of th log tstis volum-log SVL rgrssion) for M. macrorhyncha (a) and M. agilis (b) at Barra d Marica, Rio d Janiro, Brazil. Had Hight Had Lngth mals fmals mals fmals * ** 66.98*** Fig. Sb suggsts that tsts ar in a rgrssd stat during mid-dry sason months (May-July), start th siz incras in August and rmain nlargd from Sptmbr through March, shrinking thraftr. Th rlationships btn man monthly tstis volum of M. macrorhyncha and th thr nvironmntal variabls tstd r positiv in all cass. Th rgrssions of tstis volum on tmpratur (r =.93, P <.1) and on rainfall (r =.94, P <.1) r both significant, hras th rgrssion of tstis volum on fotopriod (r =.1, P =.76) as not. FAT BODY CYCLS Monthly variation in fat body mass as somhat similar btn sxs in both spcis (Fig. 6). A pattrn is mor vidntly sn in fmal M. macrorhyncha, hos fat rsrvs appar to rach a pak during th middl of th dry sason and dcras aftr August Sptmbr (Fig. 6a). n mal M. macrorhyncha, th variation in fat body mass as not significantly corrlatd to th variation in tstis volum (r =.3, F1 = 29.2, P =.88). n fmals, fat body mass as ngativly and significantly rlatd to mbryo sac diamtr (r = -.42, F1 27 S.67, ' = p < O.OS). n M. agilis, th variation in fat body mass as not significantly corrlatd ith ithr tstis volum (in mals; r = -.38, F 1 1 = 1.67, P =.226) or man mbryo sac diamtr (in fmals; r = -.4, F , P = =. 198), though th rlationship as ng ' ativ in both cass. SZ AND SXUAL DMORPHSM Man adult SVL of M. macrorhyncha (66.8 ± 4.S2 mm; n = 98) as not statistically diffrnt from that of M. agilis (66.8 ± 7.28 mm; n = 41) (ANOVA: F1 137 =., P =.997). Th monthly distribution of sizs for ach spcis is shon in Fig. 7. Mal and fmal M. macrorhyncha rangd in SVL from 36.3 to 72.2 (n = S4) and from 41.7 to 77. mm (n = S9), rspctivly. Adult mals (i.. ::::_ SS.9 mm) avragd 6S. 1 ± 3.77 mm (n = 43) in SVL, hras adult fmals (i.. ::::_ S9.9 mm) avragd 68.7 ± 4.4 mm (n = SO). For lizards ith SVLs of SS.9 mm (i.. th siz of th smallst adult mal) or largr, sxs diffrd significantly in man SVL (ANOV A: F 94 = 1.92, P = 1

6 48 C. F. D. ROCHA AND D. VRCBRADC C( 2-1 : -2 ;...J 2 t :::: /) /) -1 /) C( : ) i J C( :::: /).L a:: 2 (/) -1 /) C( : -2...J 2 c /) - 1 t : -2 8 ) i J F M A M J J A S N D ) i J F M A M J J A S O N D FG. 6. Monthly distributions of fat body mass (xprssd as th rsiduals of th log fat body mass-log SVL rgrssion) for M. macrorhyncha (fmals - uppr lft; mals - uppr right) and M. agilis (fmals - lor lft; mals - lor right) at Barra d Marica, Rio d Janiro, Brazil. al :r: f J f- 5 z 4 ::: 9 8! 3 '----.1), ith fmals raching largr sizs than mals. Sxs diffrd in th rlativ valus of HL, ML, HW and HH, ith highr valus for mals (Tabl 1 ). Mal M agilis xamind in this study avragd 66.1 ± 4.74 mm (rang mm; n = 21) in SVL and r all adult-sizd. Fmals rangd from 47.6 to 77.9 mm in SVL (n = 19), ith only to individuals smallr than 6 mm. xcluding ths juvnil-sizd individuals, avrag SVL of fmal M. agilis as 7.7 ± 5.29 mm (n = 17) and as significantly largr than that of mals (AN OVA: F, 36 = 7.87, P <.5). Sxs diffrd in th rlativ valus of all had dimnsions tstd, ith mals attaining highr valus (Tabl 2). b) :r: f- t!) _J 5 f- 4 ::: z i g 3 '---- J F M A M J J A S O N D FG. 7. Monthly distribution of SVL (in mm) of mal (opn circls) and fmal (closd circls) M. macrorhyncha (a) and M. agilis (b) at Barra d Marica, Rio d Janiro, Brazil. DSCUSSON Brood sizs of both spcis at Barra d Marica (spcially M. macrorhyncha) r rlativly small compard to othr Notropical Mabuya spcis (s Tabl 3) and to various Old World congnrs, both viviparous and oviparous (.g. Fitch, 197; Barbault, 1976; Huy & Pianka, 1977; Simbot, 198; Pattrson, 199; Flmming, 1994; Huang, 1994). Brood siz of M. agilis as comparabl to that rportd by Somma & Brooks ( 1976) for M. mabouia in th Caribban island of Dominica, but thir data ar from only svn gravid fmals. Mabuya macrorhyncha had an vn smallr and lss variabl brood siz (usually to or thr, rarly four) than its sympatric congnr, though it qualld that of a closly rlatd (and yt undscribd) spcis from north-ast Brazil (Stvaux,

7 RPRODUCTON OF SYMPATRC BRAZLAN SKNKS ) (s Tabl 3). Similarly lo valus hav bn obsrvd for allopatric populations of M macrorhyncha in a numbr of aras (both on th continnt and on islands) along th coast of Sao Paulo stat, south-ast Brazil, by Vanzolini & Rbouas-Spikr (1976) (Tabl 3). t appars that, in gnral, M macrorhyncha givs birth to fr offspring than its othr N World congnrs (xcpt for th aformntiond undscribd spcis), a fact that may hav cological implications. This spcis is strongly associatd ith ground bromliads, hich thy us as basking and foraging sits and as rfugia from prdators (Rbo;:as-Spikr, 1974; Rocha & Vrcibradic, 1996; Vrcibradic & Rocha, 1996), and has a rlativly flattnd body plan ( compard to th ground-dlling M agilis), hich prsumably facilitats its movmnt amidst bromliad lavs (Vrcibradic & Rocha, 1996). A rduction in th numbr of offspring and, consquntly, in th burdn of th brood carrid by fmals (hos body hight obviously incrass during prgnancy), may b advantagous in a lizard ith such charactristics. ndd, in th caatinga of north-ast Brazil, th rock-crvic spcialist tropidurid lizard Tropidurus (=Platynotus) smitaniatus (hich has a flattnd morphology to suit its microhabitat rquirmnts) has a rducd clutch siz (usually to) compard to th sympatric roundbodid tropidurid Tropidurus hispidus (rfrrd to as T torquatus), an xtrm habitat-gnralist, hos clutch rangs from 3 to 14 ggs (Vitt, 1981). t is intrsting to not that thr fmal M. macrorhyncha containd mbryos/ova in mor than on dvlopmntal stag (including simultanous occurrnc of stag 3 ova and stag 5 mbryos), hich may suggst that asynchronous mbryo dvlopmnt ithin a fmal may occasionally occur in that spcis (it is also possibl, hovr, that th oviductal ova in thos particular fmals had, for som rason, faild to dvlop furthr and rmaind small; in any cas, thos ova lookd normal, ith no signs of dgnration). Asynchronous dvlopmnt among mbryos ithin fmals is prviously unrportd for South Amrican Mabuya, and may b anothr pculiarity of M macrorhyncha. t is also probabl, hovr, that gg rabsorption may hav takn plac, as it has bn rportd for th closly rlatd Mabuya sp. of north-ast Brazil (Stvaux, 1993 ). W found brood siz to b significantly rlatd to fmal SVL in M macrorhyncha at Barra d Marica, hich did not occur among th populations of this spcis studid by Vanzolini & Rbo;:as-Spikr (1976) at th Sao Paulo coast. W cannot say, hovr, if this rprsnts actual diffrncs btn southrn and northrn populations of M macrorhyncha, or if othr factors such as sampl siz may b takn into account. n th cas of M agilis, on th othr hand, bliv that th absnc of a rlationship may b a rsult of th small sampl siz (th prsnc of juvnil-sizd fmals ith implantd ova is unlikly to hav affctd TABL 3. Rproductiv charactristics of som Notropical Mabuya spcis. Th lttr () dsignats insular populations. *, poold data from to or mor localitis; a, man, rang in parnthss; b, tim at hich parturition occurs; c, calculatd from Tabl 4 of th rfrncd papr; d, as M. mabouia;, as M. bistriata. Spcis n Brood Siz' Rproductiv Locality Sourc sasonb M. agilis (2-6) Oct-Nov Marica, S Brazil Prsnt study M. bistriata 5 - ( 4-8)*? Amazonian Brazil Avila-Pirs ( 1995) M. caissara (2-8) Nov-Dc Ubatuba, S Brazil Vanzolini &Rbo;:as-Spikr ( 1976) M. caissara (3-9) Nov-Dc Caraguatatuba, S Brazil bid. M. caissara 1 4. (2-6) Nov-Dc Sao Sbastiao, S Brazil bid. M. caissara (3-6) Nov-Dc Brtioga, S Brazil bid. M. frnata (1-8)? Araguaia, Cnt. Brazil Vitt (1991) M. frnata (2-8) Aug-Nov Valinhos, S Brazil Vrcibradic & Rocha (1998) M. hathi (2-9) Spt-Nov xu, N Brazil Vitt & Blackburn (1983) M. mabouia () (3-5)? Dominica, Wst ndis Somma & Brooks (1976) M. macrorhyncha 2.4 (2-5) Dc-Fb nsada, S Brazil Vanzolini &Rbo;as-Spikr (1976) M. macrorhyncha (1-6) c Dc-Fb Pruib, S Brazil bid. M. macrorhyncha () (2-4)? Buzios, S Brazil bid. M. macrorhyncha () (1-4)? Qu. Grand, S Brazil bid. M. macrorhyncha (2-4) Nov-Dc Marica, S Brazil Prsnt study M. nigropunctatad - (3-7) Aug-Nov lquitos, Pru Dixon & Soini ( 1975) M. nigropunctatad ( 4-6)?Mar-Aug Santa Ccilia, cuador Dullman ( 1978) M. nigropunctata (2-9)* Aug-Spt Amazonian Brazil Vitt & Blackburn ( 1991) M. unimarginata (2-7)? Costa Rica (Pacific slop) Fitch ( 1985) M. sp (1-4) Jan-Fb Cabaciras, N Brazil Stvaux ( 1993)

8 5 C. F. D. ROCHA AND D. VRCBRADC th corrlation, sinc only on fmal smallr than 6 mm as prsnt in our sampl of "gravids", and it had only thr ova). Brood siz is significantly affctd by fmal body siz in othr Brazilian Mabuya spcis (Vanzolini & Rbou9as-Spikr,1976; Vitt, 1991; Vitt & Blackbum, 1983, 1991; Stvaux, 1993; Vrcibradic & Rocha, 1998), and it is quit surprising that, in our study, M. agilis did not sho such a rlationship, hras M. macrorhyncha, ith thir smallr and lss variabl broods, did. Lik othr Brazilian Mabuya spcis hos rproduction has bn rasonably ll-studid (Vitt & Blackbum, 1983, 1991; Vrcibradic & Rocha, 1998), M. agilis attains rproductiv maturity at small body sizs (i.. about 49 mm SVL), prsumably hn only a f months old (s Blackbum & Vitt, 1992). Mabuya macrorhyncha, on th othr hand, apparntly dos not rproduc in its first yar, as suggstd by our data and by Vanzolini & Rbou9as-Spikr (1976). Th lattr authors also mntiond that th smallst rproductiv fmals in thir sampls r about 6 mm in SVL, hich agrs ith our data. Similar pattrns hav bn rportd by Stvaux (1993) for th closly rlatd Mabuya sp. This rlativly dlayd rproduction of th M. macrorhyncha linag rlativ to othr congnrs (including th sympatric M. agilis) is difficult to intrprt in th light of our data and dsrvs furthr study. Nvrthlss, th gstation priods of both M. macrorhyncha and M. agilis ar apparntly idntical, spanning btn nin and tlv months. Th pattrn of mbryonic groth is also apparntly similar btn th to spcis, ith littl incras in ovum diamtr during th first fiv or six months, follod by rapid mbryonic groth thraftr, as in othr Brazilian Mabuya spcis (Blackbum & Vitt, 1992; Stvaux, 1993; Vrcibradic & Rocha, 1998). Th rproductiv cycl of M. macrorhyncha appars to lag about on month bhind that of M. agilis: of th 17 gravid fmals of th formr spcis collctd in Sptmbr, all r in stag 5, hras fiv of th ight gravid M. agilis from th sam month r in stag 6 (i.. bor llformd mbryos). Thus, M. macrorhyncha brds somhat latr than M. agilis, ith parturition probably bginning in lat Octobr or arly Novmbr and apparntly xtnding into Dcmbr, hn that of M. agilis may hav alrady casd. Unfortunatly, hav vry f adult fmals of both spcis from th priod Novmbr-Fbruary, so that it is not possibl to dtrmin hn parturition actually cass in ach of thm. t is also intrsting to not that th thr smallst gravid M. agilis from Sptmbr r in stag 5, hil th fiv largst r in stag 6, suggsting that firstyar fmals of this spcis may brd somhat latr than oldr fmals, as rportd by Blackbum, Vitt & Buchat (1984) for M. hathi. Although lat-stag mbryos r found in six M. agilis and to M. macrorhyncha, non of ths appard larg nough to b full-trm. Among Sao Paulo populations, nonat M. macrorhyncha ar apparntly born at a SVL of mm (Vanzolini & Rbou9as Spikr, 1976; Zanotti t al., 1997). Thr trm mbryos takn from a fmal M. agilis from th rstinga of Grumari, locatd about 16 km st of Barra d Marica, avragd 3.6 ±.38 mm in SVL (Vrcibradic, unpubl. data), hich suggsts that th young of this spcis ar born at a SVL of at last 31 mm. Th lack of nonat-sizd individuals ofboth spcis in our sampl furthr obscurs our undrstanding of hn parturition actually occurs. Th brding priods of M. agilis and M. macrorhyncha ar short and ll dfind, lik thos of othr Notropical Mabuya (s tabl 3), hich ould supposdly plac thm into th catgory of "non-continuous" brdrs, according to th classification of Shrbrook (1975). ndd, basd on that ork, Rocha (1994) rfrrd to M. hat hi and M. nigropunctata ( = M. bistriata) as having non-continuous rproduction. Although tstis cycls in mals of th Barra d Marica spcis ar clarly sasonal and non-continuous, application of such trms to fmal cycls may not b appropriat: th simultanous prsnc of vitllognic follicls and implantd ova or mbryos in th spcis studid by us indicat that rproduction may actually b continuous, ith ovulation occurring shortly or immdiatly aftr parturition (almost all sxually matur fmals of both spcis r rproductivly activ). Th production of tiny, yolk-poor follicls by fmal notropical Mabuya is nrgtically unxpnsiv, and may occur simultanously ith gstation, hich is vry long and accounts for thos lizards having annual rproduction. Mals of th to Mabuya spcis, unlik fmals, apparntly attain sxual maturity at similar SVLs (55-56 mm), although th minimum rproductiv siz of mal M. agilis may b ovrstimatd, sinc it rprsntd th smallst mal in th hol sampl. Tstis cycls ovrlap considrably btn th to spcis, ith maximum gonadal activity during th t sason, coincidnt ith th priod of parturition and ovulation in fmals. t appars that, for som rason, tsts of M. agilis suff r a gratr rduction in siz during th dry sason compard to M. macrorhyncha. Rproductiv cycls (both of mals and fmals) of th to spcis ovrlap almost compltly, hich suggsts that thy may b rgulatd by th sam factors, such as nvironmntal cus and/or food availability (s Rocha, 1992 and Stvaux, 1993). nvironmntal variabls such as rainfall, tmpratur and photopriod sm to strongly influnc th tstis cycl of M. macrorhyncha, hos rspons to th variation of th first to apparntly taks about to months (it is quit puzzling that th rspons to photopriod as diffrnt). Mals of anothr Brazilian spcis, M. frnata, also rspond significantly to th abov variabls ith a tim-lag of to months, but th rlationship is ngativ in this inland form (Vrcibradic & Rocha, 1998). Similarly, tstis siz of mal M. hathi from north-ast Brazil incrass as th dry sason progrsss and dcrass hn conditions

9 RPRODUCTON OF SYMPATRC BRAZLAN SKNKS 51 gt ttr (Vitt & Blackburn, 1983), shoing a trnd opposit to that of M macrorhyncha (and of its clos rlativ, Mabuya sp.; s Stvaux, 1993). Mayb th mal cycls of notropical Mabuya ar mor strongly tid to th fmal cycls than to dirct xtrnal influncs (mal pak sprmiognsis alays coincids ith fmal lat parturition-arly ovulation priods), but a bttr undrstanding of th ffcts of nvironmntal cus on mal cycls of tropical lizards is ndd bfor any conclusions can b dran. Monthly variation in fat body mass dos not sm to b too important for mal rproductiv activity in ithr spcis, although a comparison of Figs. 5b and 6 (lor right) suggst an invrs rlationship btn tstis and fat body cycls in M agilis that could, prhaps, b clarr ifth sampl sizs r largr. A comparison of Figs. 4b and 6 (lor lft) is vn mor suggstiv and, again, bliv that th small sampl siz of M agilis as rsponsibl for th lack of a significant corrlation btn fat body mass and mbryo siz. This as not th cas for M macrorhyncha, for hich th corrlation as significant and shod that fat rsrvs undrgo a dcras in mass during th rapid groth phas of th mbryos, as in othr Brazilian Mabuya (s Blackburn & Vitt, 1992; Stvaux, 1993; Vrcibradic & Rocha, 1998). Th considrabl incras in siz of th concptus of M macrorhyncha, hich is similar to that of othr N World congnrs (Vitt & Blackburn, 1983, 1991; Blackburn & Vitt, 1992; Stvaux, 1993; Vrcibradic & Rocha, 1998), indicats that rproduction is mor xpnsiv nrgtically for fmals than for mals, hich may xplain th diffrntial importanc of fat rsrvs among sxs (.g. Gaffny & Fitzpatrick, 1973; Jamson, 1974; Ortga, 1986; Ramirz-Pinilla, 1991; Huang, 1997). Th pattrn of sxual dimorphism in siz and shap obsrvd for th to spcis at Barra d Marica, ith fmals raching largr body sizs, but having rlativly smallr had dimnsions than mals, agrs ith that of othr South Amrican Mabuya spcis (Rbour;:as-Spikr, 1974; Vitt & Blackburn, 1983, 1991; Stvaux, 1993; Vrcibradic & Rocha, 1998). Sxual dimorphism in M macrorhyncha is also clar from Tabls 6 to 11 of Rbour;:as-Spikr (1974 ), concrning populations from th Sao Paulo coast. Larg fmal body siz among notropical Mabuya may b th product of volutionary prssurs acting to incras th numbr and volum of offspring carrid by th fmals (.g. Fitch, 1981; Vitt & Blackburn, 1983, 1991 ). Such a prssur ould not b xpctd to b strong in M. macrorhyncha, hos broods ar usually of only to or thr (s Vitt (1981) for data on th smallbroodd Tropidurid Tropidurus smitaniatus),but vn so, fmals of this spcis hav SVLs largr than thos of mals and positivly corrlatd to brood siz. t is possibl that small brood siz in M. macrorhyncha is a drivd charactr, acquird aftr its ancstors adaptd to a bromlicolous mod of lif (s abov), a vi opposd to that of Rbour;:as-Spikr ( 1974), ho proposd that this spcis is an ancstral form, basd on biogographical analyss. Naturally, biochmical and gntic comparisons ith othr N World spcis ar ndd bfor any conclusions can b dran. Among mal notropical Mabuya, attainmnt of rlativly larg hads has bn suggstd to b rlatd to aggrssiv mal-mal intractions (Vitt & Blackburn, 1983; 1991 ). This sms a plausibl xplanation, though hav nvr itnssd such bhavior in ithr spcis at Barra d Marica (s also Stvaux, 1993 for a discussion of this topic). W conclud that M macrorhyncha and M agilis hav rproductiv charactristics and pattrns of sxual dimorphism that ar typical of N World Mabuya in gnral,.g. ovulation of minut and yolk-poor ova, gstation lasting about on yar (ith most of th mass incras of th mbryos occurring ithin th four months prior to parturition), larg fmal body siz and rlativly larg had dimnsions in mals. Mabuya macrorhyncha has, hovr, som pculiar charactristics that diffr from M. agilis and its othr Notropical congnrs (xcpt a vry clos rlativ in north-ast Brazil): it dos not brd in its first yar (dlaying first rproduction until attaining a SVL of a. 6 mm), producs rlativly small broods, and fmals may prsnt asynchronous brding (or ngag in gg rabsorption). Ths charactristics should not b attributd to possibl ffcts of th sympatry ith M agilis, sinc thy ar prsnt in allopatric populations in Sao Paulo Stat (Vanzolini & Rbour;:as-Spikr, 1976), nor to th fmal siz-brood siz rlationship, sinc th to spcis do not diffr in adult body siz. Such uniqu faturs of th M macrorhyncha linag should b bttr studid in ordr to invstigat if thy rprsnt primitiv traits or scondary adaptations, possibly rlatd to th spcis' bromlicolous habits. ACKNOWLDGMNTS This study is a portion of th rsults of th "Programa d cologia, Consrvar;:ao Manjo d cossistmas do Sudst Brasiliro" and of th Southastrn Brazilian Vrtbrat cology Projct (Laboratory of Vrtbrat cology), both of th Stor d cologia, nstituto d Biologia, Univrsidad do stado do Rio d Janiro. W thank P. Tixira-Filho, S. Ribas, L. F. da Fonsca, L. N. Martins, A. M. da Silva and M. Cunha-Barros for hlping us to collct th 1 izards. Jorg P. das Nvs prpard th slids for microscopical xamination of lizard tsts. Jams R. Dixon and Moniqu Van Sluys kindly rvid th manuscript and offrd valuabl suggstions. Th Sub Ritoria d P6s-Graduar;:ao Psquisa of th Univrsidad do stado do Rio d Janiro mad many facilitis availabl. This study as partially supportd by grants from th Conslho Nacional do Dsnvolvimnto Cintifico Tcnol6gico - CNPQ (procsss N 43787/ 91-2 and N 3819/ 94-3 NV), and from Fundar;:ao d Amparo a Psquisa do stado do Rio d Janiro (FAPRJ: procss no. -26/J 7.385/97) to C. F. D. Rocha.

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