03 vial bolitoglossa

8/18/2019 03 Vial Bolitoglossa

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R BioI Trop, 15(): ,·5 96

The Ecoloy of the Tropial Saaand, Bgs

subm in Cota Rica

by

L. Vial'

INTRODUCTION

BACKGROUND AND SCOPE OF WOR

ode ecological oncep have be etabihed ad refied moty frmda obtined i north temprate zoe tudie. Dpite the fat that about nethird of the land masS of the earth ad ary two-third of it biota ocu! in thetropi one, litte j known about bai biological procee or ecoogia retion

hip der the inuee oE the tropia un Te tropia enironmet i ditinguihabe from that of extatropial atitue on the bai of photoperiodiity,temperate, heat budget ad prodtiiy; a combiation of factor whihprouce uique eoogic ettig

Atough umerou work hae deat with he tropi iariaby the defin·ton of the regi ha bee iadequate from an ecologica poit of iew In orderto ealuate the haracteritic of tropial and temperate regio it i neeary topreent a foudto of ommon undertandig on the ature of th tropical eniroment

he tropi are uualy defied a the wm atitude yig betwee 23' N ad of the equator Beaue of the ' reatio to the eath thi latitudia rego reeie maximum olar radiaion f he earth ufae were o uiform ompoitio and phyiography it might e epeted that a characteritieiromet inhabited by a topica bioa would be foud ro]ghout he regiohe nouniform ditribution of land ad water and phyiographi ierity of and

mae modify ditributio o he heat budget o hat irreguaritie ocu On thebai of ow imit of ditributio of tropical biota it ha been uggeted thahe tropia enirome i it broadet ee i limited on the north ad ouh

Dearmnt of log Unjvrsiy o Missur Kansas Ciy Missui.

1


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REVSTA BlOI PCL

n eestrl nd reh water uations by the 4° e man nl them oreed o sa ve. rpi mrne abits ocr in are we men u·ace tmpetue n h codes monh s grter han 210 C The tropca regini here defined a therml zon occrrng n exemes 5° N L and abou 3°S at Vats in nrh and suth extent of the tropic boundr re du tosie nd levton of he includd ld areas and effe of oceani curn

Withn hs rgo the daylgh period s neve! ess han hou in 2nor mOe han 14. Farthr north nd �outh the photpericd my rnge m eroto 24 hous dependng upon the on Solar [diaon in the opíc bel averges 860 gm cacm'/dy range 720 o 1,000), wher in exa toal ras the

vage s 60 gm cl/cm/day ange O to 1,00) Cen per comm ahe budget ba he to receve he gett pprtion oE sr egy ath energ i5 evenl dbuted ovr he ea

Lk of seaonl s omt regrded s a ropicl hcter, bt callhe on ae obred beaue dl xtrems E tmperre nd hudity so greatr rnge of arian han nnul lmac c e Saona changs in he - pics are expesed incip b dffernces in ranfall ditrbuon

n a gnel way there is correspondence {f bota wih the deli tngtherml boundaes alhough broad areas of tnsition m b peen Hgh mon-tan and alpne regons dct much of the ropi. \th the 24' C e vel)soherm thee go are topical n hat hey a expoed to he sam lght perid-ki nd energy magnds a the ownds alough their matíc parmeer ma a rt igh corrpod o he f etratropc lcaltes

xtratropicl plhodonti amanders h ben n of th mot widstuded group o mphbins. he gea dversity whn h amy nd s boddstrbuion of exellnt opporunies for sdies of compaatve ecog ahve been dmonrated by DUNN 2 GOBMN (32), HTON (3, dGA (56. Although he Pthoontidae s h only famly of salamandersfound in he tp, o neive sdy o opicl repreenttve has be

prvously attempte rbb ew oter speces of salmands are so st foutned fild esgaion a s Eo/ilgu shpamf (Bouenger) AI-tough praccal ojng ws own o its ife hsty he p remabebundanc on th Cero d la uere of Costa Rica indicatd an almot idltion for cntinuo ud thghut i vrcal rnge of dsribuon accessbe by e ¡temrca Highay (Fg 1.

Th obecve of h rsen tudy ae vefold: (1) o detrmin heloca disribution habita eleion and life

hry of B bp; 2) o

evaluate the gifiance o ouion chaes s eresed b the cologyo he pece; 3 to provde unde"tandng o th bú boog oalmdes in he rpic wth ptla mhsis ecologic difference may st betwen pouns n top and empera znes (4 to determhe fesibty of ppying ecoogi and demogahc mehods to topiorgnsm and (5) to devlop foundion for fre work in tropcal poplnecoogy.

Pemny d wo ws crre ut in Ju o 96 Fultme tudy a


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, fCOOGY O OTOGOS SLMT N COT RI 15

bgun Aust 196, nd otiud tí! S


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ECOLOGY OP BOITOGOSA PAMAA N OA R 1

ama \ee en retued t the spt at whc hey "'ee taen Alhougmarking f new indídual was pfmed oly wtin a dfed area the surodi 0 m 00 ft) zone was eaed in the same manne maredsalmandes ! eaptued amas wee studied bve. the sites 9idivda wee marked d st

died dng he yea

tudy aeas aed Erom q. m (700 q t.) t 0 sq m(18 s Et The size oE eah aea wa detemined pmaly by the numbeo ams tat oud be psed in a sne day tus aidin any nona

poplati sEt

Meauemet E ditans ee made with a 0

m 100 t.) Ln stetap Azmt adins to te est degee wee take wt a Uüed Stats Ay

enatc cmpas Cae was praticed t ad any mtic attractin by etaneO objec

Na at o B. was stded on dffent ocaon the eo de la uete betwe ,60 m 00 Et) and te summit Pduk t podawn bevatos were made at five ocaite whi nuded temaked popaons I ah sdy he site was cheked ng a battey eadamp at two ou interva thougt te niht. A consstet pah f eaminato f te aea wa fowed and eqa time alltted t each vit Follow themod oE N () a salamnde wa condeed 00% ative E it adompetely emeed m cver and wa fund te pe or 0 ae btpartially exposed To avod egative espne by ative mals noe wee touhed o distubed n the nigt f Ma 2 9 at 9 m 900 Et) thesouten ope a aive spmen ee ceed at eah st Afte ne heaea wa toohy ·examied ad al aailabe samades wee colleted.

At te begini and nd of each viit elatie hmid: nd o th michabtt soil and ai wee reordd

a mnth amples E B. bp wee andmy take at attudial

ntervals oE about 0 m 00 Et. ) ; uadat ste ad maked poplts weaúdd AH pecimens were nted as tú tandd lngh nbe and sie f testula Jobes O numbe ad sze oE va a wl as the coítion of oday sechaactes. Frm eac ample paaff ectn were made frm indvidual ected at mm ize terals t determne te rlationship of ie and s�ual deveopmet Gads rom saamades wee Eed y oution tined witematln-eoin ad sectoned at 8 to 10 micra

ggs and attending aduts obsecved in he Eield wece exmnd at intervalsand rd takn o eg zes and stage E deveopment rten eg ut

wh the .tendig adJts ",er bought to the labratocy and mantaned at on·din appximating the f the naal abitat i de to pemt study ·Ebdin bhvio and dveopmet. Iormatn wa obtned Erm a ttal oE 31cltches d atedin aduJts.

To oberve courtsp and breeding behavi males d females we kepttoether in captty. Te aims were eplaed at bi-weely inteals nureoptima pysogca cnitn Twenty-ou expementa aims wee ub


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18 V DE

anouy mpaned w puay gads om s Ra piPem o B Ib-

lJa o stmuat dig nd ovipto.aboratoy pments wIe cnducd o tempatur toeras ad water

rqumnts of B. t as wel as dhydrato ad ehydran ratesCand sil sampls w ake n th fild durig most peods of udy temakd populaos, quadats and aciy cyles; clasifid as textue ad clh wighd befr and afr heang at 15' C fr 48 hOs oil oi valu,re prssd as pertage f dry wegh

AH measurmts spcmes ar n millimtrs Ule onvi �ad,ss o dvduas ae gve as standard legh (nou t pGserio magn vct).

Masrments on liig spiens wee made by ung a ged b

ard abu cm n lngth wappd i abrben pap oweig e l was plaed i

he grv drsal sde wn and a ll u pacd genly bu fimly r n sufce Readg we e o e 0.5 m epad measmes o e same anm jnded an acacy o ma�emnt o ± mmn adts. Safced oms w ked n ydos obutano (Croead mard with a els dl alp to th aest 0 mm bee esati Rpeated measurements idcaed an acuracy of 2 mm.

M

Ths prt s a ou of c ctibut o eOS pe adsevral agcs. y iebtedess o the Unierdad de Cota Rca ant emagnfid h entr searc pogram was accomplshed durng y affliatwih uvrsity f as Vsting Professor f Zolgy and later as Fd Fnd-í Prfso of Eology n fe yeas 96 hrugh 1964 Durg hi mo-abl perod my wr ws thuiastally supprt by Ds. Rafael Ls Rdíguez ad John eAbae Prfesor Eot Ce Chef of t Seco Meo-ógco d Csa Rca mde aaab s ogna da o opa mas ad -

ssd in coeon o w ods, as dd S. Vcne Conto PdoVo A ored rquen tecic asssane d Sa Esa Poa peromda1 mcotchnica laos. S ame Vla R ad with seeral f he drwns.ng Fdrico utérrez B Drct of the nsttto Ggráfic de �_t aovdd muh of th opogrp ifomai

r ay M avage, f he Uve"y of Suthe Cafria suggested thrgina prblem an hs ontiud gida peased to akwe DsRobrt M Chw and oh L Moh ffered numO sggetins n hei t!evews of th dissrtaion manuscpts, fm which this pa has b exat

and rvisdhe help and enouageme of my fe, Lynda W Vial tbue g:-y to al phass f h wor aso wsh to exprss my graiude o may ndid-uals who by hir anonymus asane n s way ntrbued to sdy

Fnancial suppot was gntd by h Aman Phlosph oey eos Fun Gn N 3029) th Naton Sen Foundaon GE 4705) an Fod Fodao awad vsdad d Cos a Te Sc o Gad


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SS N S RIC 9

Suds, Vvit f Mu a Kna Ciy, gnrusy ubbed he sof publicon.

THE ENVIRONENT

lOCAON ND PHYSIOGRAHY

The Cdir mca dmne h gry f so CoR. t th norhrn magn f thi ang H h Cr d Mur 9 5'N Lat. 8 5 W Log ) n f e argr mfs, og n vi of349 m (110 . nd uuoded by seve saUt aks f imlar hghtt i hrgh nd o s rgon that h te·Amra Hghv

y me$ nou

ri tran rahog it hght eve at 327 m 069 f ) Th dyegio (Fg ded a ar! o s hghway aec soe k lgh ada lied eat ab 1 n ithe ide E th mmit Anamit wr tabid fr drming the a dtibuion f B bpd wee as 150 m bw eve whe s slmode w cy fOd. Thvertica rang from h owst i h uy aa h mm i abut1,100 m.

xteodig in a N·SE cn fo r 75 m h Crr a·

mana ¡nud era k f nay 4000 m forng r nifm ntie i whih h w (Paic) d m abrpt tha t eate tla) Pimry grtc i h be inrrpted by oca onatiity nd ompex gec pese dung ri f rgn whihda rm edy Moene LOYD 5 ; \EY 79 80). Cr Mrt h gh regn in t ira acceib by oad In yar pk ha bn rnamd Ceo Bn it thugh the mif ít i i erdo by he eer m

Si h "pp p f h ori a pry T

h

iz

s

a ay s th a e i dph Freqeúy a i um í n ott wth ighty wahr rc mate. Du tú h me ogc stur i xr nd lor ae aie At tin t appxmat2,35 m, graybow ad ad omy doiae a1hgh bk m ay r in itd ar. Cyy o uay ywgay a ommon bteeo 240 ad 2,75 m At high vatos gay·brw gry a ia thg htrd aa ad deios dpoits of bak siy 10 u; resut f ozd min f gio WR 8 pot au f 9 t 0 f am fom Jboni ak

CLMATE AN WE

e n'rrw f tlC hmu t th attd (10 N) whi ah Pai On an Carbban by than 10 ri Ca Rawh n ma ang ny gnicnt onnnl fne Wtn gene an toca at wr many a [at h a t-gaphy, i rn n t ak t tring dparitie


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EOlQGY F lTlO SUBMA IN RI 1ABL

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22 VISTA DE BIÓLGt OPA

oon afte noo) foowed by heavy but sporadc ra In late afternoon so therm

! tabty is reched and e ods gen Vb altrnate ro r tovera! hndrd metr winds ove h 10ud ovr ountain ridg Drns oowd by spotty oud cove but uring th ngt there is gadu arngo the , a reduton o humdity and decree in temeratur

VGAN

The area nuds two mj matc egetatio formatios: the Topalower Monan Wt oret and te ropical Montan Forest (HDRICE,

40 4 In thi part o te ordera thy have uppr i at 25 m nd3 , respetivey Aove the latter elvato occur poory dned dis-clx couny he áo reulng from e no o e burned ovrsit regin y emnts f ubapne wet woodlnd It c howeer imatc oraton, as the temperatr and prepitton re charaeristic o theMotae Rain Forest Botemperaturs d vegtatonal ortons ssoated tho! atc ater ae ncded n Tbe

Ech oraton ractzd by cean plant ecies whh reet somdegree of doiane and rrton distribution Nomenlatur and pecesdeterminton based upon nformaton r H ( NY 68) ,

and 78.On the Cerro d a Muert dminant tre pee of the Lower Montne

et orest inude CO"n!I difla, Cdrl Todz, AlnJ jo gnla poaana Dmy gandeJ and Per Schedan Seera peces eah oíMn óega and Pipe ar aundat n th woody ndergrowth. O pooro Qers ps or Q oocr my or s most pure tands reachngheghts o over 3 m he genera apet of ts formation is shown in Fig 6

In te Montane Ra Forst Bde pin Hpmle obov, El-01i pa úía pnaü {úa ata, Ddmpax i d oo-

c'p mont are charatestic tees speces. The arge ok Qe Ocen, for nse extensve stads domnaig the physogoy of m o hioaon Above 3 ee) coea, Vm ag1 ¡pem rtm and H iode re o of the mor ommn hrus wihthe conspcuous dwarf bamboo Chwq a, reprent a mitre omontan ad pámo spees (Fig 7

n te otane ort trnks and rnhs of te rger tre are saUyhdden der a covr of epphytes Ptrdophyte, broids, orchds a roidscompre a majr part o these "aeral wap ommuniti. he groud s co\'e-

d by nmerou dmbent herbs, mosses nd lns aong wh e see oAlhemila, Gomoz, Lopdm and phgm n oal swampy depressionsthe tre rn, oaí o nd th lge trrstral bromeld, Py dayode emrge rm a dnse mat o sphagum os to presnt a alot eeresecale (Fg. 8 . Oe d dbd ae oed b g c-na Cir bo'en Rm oce nd ptar largel_d G¡e Ig d G. cana


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ECOtOY F BOlTOGLOSSA UBPM OSA RrCA 23

RT HTT

Bg bpma cuies exene ang jn e empa eations o ower Centra Ameica It as be epoed om he Codie ne ppe margis of he Mee enal a e Codi d Talma Cosa Rca nd fom egon oE Cap Boee Pam n e lor lisof its atitdna age, ppomly 10 m hs sam s encousporadcy ad s comm ony boe 1>5 m he secs s hactertc of teoica !ower montane d montane egons n a geoahic nge bsced inortern and outher prs by e Mesea Cenr o Cota Rca

On the Ceo de la Merte B inhabts a bd atudi beabo 2,300

on he nrwesen soes, desies concu�ly ¡crasig

with elevaion O he suther soe is a but absent beow 2,00 mBy comarg inomaion rom pecpai a temeaturs, a obios

correation o pouao densHies wih matic conions is see. I hs begney accted hat mosture as eprssed in e amount and isribuion oana eets the geet infuence on he dstribuion o Qrgaiss n oicalenroments· (L RU 49; HARDS ) and ha emeaur i secondary mor ce hi is ony e o regios with faily unfom surce aMany aeas o low eleatons, .., below 1 m in Cosa Rica meet e mois-

ture d rfug requiemnts of B Jbpm ven gea rainfaU is chaaceiti ceain coastal regos of he couny, ye owland oica we foes aeasa no ocied by hs secies. Apaety tmpeae is he domnant huence resticng the disbuo o hse samandes No miting bio-ogica fao c yt be suggested The seces has no been obseed to ocupyhabt on b Cerro d Muee a bioemeatures ighe than 1 5.2' C 1 doo imply a micoclmae of abias s measuable by cods fom weaesaions, bu ceny mcaes ae nflue in a geeal way by gos mat rgimens IE 7)

H TWiin s altidal ange, B bpm occpies dese abas

e Topial Mnae Rn Foest, wee empeaes ad evapoansatíolim pos no pbem e sallow cove povde by e and os s common eea Te Ceo s sbec o sfae ing dig wine b s-amande popaions ed miga only a few cenimeérs elow e gons;fac in cevices ad roofactued canies avoid desÍco.

In midle elevons ougly beween m and m e seJcedabiats ae quie difen. he dense cpe of spagnm d mos a

eans a hg moisue ee ad a sabe, eavey w emperatu is a hbamost reueny uized. Occasony saamades may be encoueed underoks and pl debris b only when ese ae deeply mbeddd n he soil oposse a wedeeloed ichenmoss mag Whee ocl se bank opograyo sistent ras ody lcal mose condions habtats ht would oter-wise be unaorable can be ocpied empoarly.


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VIAL: ECOY F BUTGL0SSA SPlMAT JN CS RI 25

ng te spermatoyte 1 phe. Eah of ese obes undergoes maaton as a

unt th ne lbules beng cntnually addd anteroy.As a esut of ths pattern of formaon, maure sperm ft appea in t

mst posterior part o the lbe nd grduy develop mre anterioy as adjaentbles mae n sequene The spermatogen ave" s actualy a sees of phaesn the meotic proces each of which is confined to a snge obule.

A fairly constant patern of tnsion of gem es can be observed wthinthe tests of B. mbpawdü, (see Fgs 9 and 10). he transitonal stages frspermtogoni I to speatoytes 1 usualy ocupy the greatest par! of the tstsexcept for temina primay lobes in whh matrton i the anteror poton has

been ompted Te region ocupied by lobules ontaining spermatoytes II srelatvly narw, ndcang a apd transon from spermatoytes 1 to smadsLbes ompsed of permatds and spermatozoa usually oupy a regon abouteual to tha! of spematoytes 1. This impies that spematocytes and spermatidsrequire a longe erod of matuatn and tansfomaton tha the mitotc dvisiono sermatocyte H.

As the testis bees fully developed moe lobues prduce spermatozahih are iberated to the centra du In soe aimas al obules may be n anadvaned stage After lbron f sperm the oules degenerate and atophy helobe the ontinuay moves foward as the primary germ ce ord s ativted

In a sense the lob matures itself iota non-estence Te poterior dgeneratedrgan beomes tbe caudal gem ell od o tempray sterl" se. hs egn'hh is he basis of he lbed naure of the rgan, wil evay become retivated and gve rise to spmatogona and the interstita sepa of the secondaryand tertay obes n the se manner of deeopment a the prry Uslly eposteror obe arses befe the complete mauraion f the bes n the anteorne. Advaned stages of redution dvision are not always present in the seondand thrd obes o testes dependng pn the sze o the lbes; 10r s it euayresent in the rght and left tests f a sge anmal

REPRODCTVE CYCLE O FEMES

Oa are frt dstngishabe when abot 0.1 mm in daeter As matu�aion otin1es they gradally incease to an obseved mxmu diameter oE 3.5 m befre ovpostion. Wih e increased acU ulatn of ylk, the egg bomes cem-whte in col Defntely, not a E the ova matue. In alier stages the1mber vsble ranges fom 19 to 7 1 i a single oay, t duing the later sagesa ize differental bome onspcuous as soe ither acumulae yk nor sgnifcany enarg The average nmber of apparenty ·'úp" ova (> 2.5 m dia-meter) fund in a single vay was 1 5.2 ( ± 1. 3 stndard erof) n a sample of 40 femals. By omparson, e arage nmbr Df es (2.6) bserved in 3 1 cJtche indiates that both ovais mature and ovate duing th beeding eioBeaue oE the great number of mnue Ova pre:ent in cay devlopment, it has not been feasble to determne what proporticn a.Iy maure and are ad; how.ve, s a eltive y small at oí th tota that aqus any qua o yok


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ad pass ito he oviduct durg oviposion So indicaton tha immatur ova

are reaned ad may subsequently maure is offered by v bodg fmalesthat cotad numerus small o meum (0 to 20 mm da!e) 'hs ovain h ovares.

he oviducs o maue dividuas ar eargd, thined hghy co·volutd tubes whch occupy much o he abdomal cavy immatue spmenhey ae thn ad taght dorsolaral bules

Feaes o B. b(¡M do o reproduce ve yea Idívdua$ ihenlaged ova were ud at aH ms o th yea bu mos o th marked adultsdid become gravd dug the perd of study wo maked femals rmad gravid over a perid of eght months wihou ay nrvenng oviposito Un-

forunaely, o informao i availabl on marked gravid anmas ha subequnt-y were Doed as "spn The female elme o he populaio is capable breedig hughou the year Ahogh n ceraL moths a greatr nubr ofravid mals wa: observed, he actual proprto total sampl siz di o oably de

OVPOSTION

The sge occason o egg laying observd i B bmt occurred i a

ad fmale whch had b subjecd o a subcutaous gula mplan of a sngeaerior hypophyseal obe of esh R"na ppn pituitary on May 9, 962 Arimplanao, the salamander was kpt mst file papr a coverd, 10 cmcultue dh paced a darked rom a 0 to 22° C.

Approxmately 7 hous lae ovipostion bga wth he extrusio o age egg; wthin the ext few miutes two more wr delvrd. Th feaJspasmodically archd her ta just bhnd the cloaca ad lted he pelvis by xenson of the hid lmbs before ovposto Erac spasms orcd the eggs downth oviduct to the cloaca ad relaxao of th sphcr prmted extruson f

h oa he sngly or seres. Upon mrgence of h las gg, i was "pinchdoff by coracton of the cloacal lisurg a period of our ad oe-hal hous he fmal laid ght eggs

ad a tervals movd reey abou h moist contai Sh he colleced theggs nto a small lutc by us of her al T hours a he las was xtudd shded, probby om hormoal shock. A his tme ou udliveed eggs wevisible h abdomal cavy

NSTG SITS

Eggs of B Sbm" are dpoited undr wimbdded rocks or dayig ogs n hese atural depsios no excavai or modiicao o the t appare h se of he stig si neds oly to be lage ough t acmoda he pae ad egg cluth Th sie s usualy locaed wher ther ha beno physcal distubace or �veral yars sic the surface s coveed wh a maof ell-developed lichn and mos colonies Occasionaly nsing sit a und


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28 REVIS E ILGI RPIC

t a mnimm and obertiS made a nfrequently a but ne every to weeks,

the broodng adlt 'nvariably abandned the uth Flng t, he eeaed deveopment and dempedWhen found, four of the egg luthe ere .tended by mae Thrteen

he ere mantaned n the aboratry th the acompanyng ault, ne Eh a a male No ditintve feare ere derned n the brdng be�havor f the ee

he aending arent uaHy a oled tightly abot the egg luth ththe frelmb and throat regin reting n the upper rfae f the ma hembrae a ualy utained for three o fr day tht ntabe hange Durng the perod of brding the adut edm left the egg and a alm! ayn onta i them At interva of evera day the adlt ere oberved to rtatethe uth th the forelmb and tail, bt hangng it latn tte nt at aH

videne ím boh abratry and fied tde ndate tat pertentbt not ontant attendane E the egg by an adult a rete f their ntinued developmnt under natral onditin In aptviy adlt ere berved toeave the th n oain and to ander abt the ntainer premaby foraging for fd (he digetive trat of three frehly apred broodng femalentained part of net) a of ntant attendane i mt nvnnglyeablihed by the preene of a mal in the abene of a female

EMBGN

No eavage i vbe n nly depoted egg The earlet tage reogniz:ble ithut arfe of the egg that f the era grve Opt vele and eappear oon after onvergene of the neural fld and nderg rapd enlargement, bemng the mt pu fture of the pot_eural embyo Anterorlmb bd moth par ad gill flament aring a 1tgroth frm the garfod region appear at apprxmaey the me time; the poterr imb bd

beng apparent omehat ater By the time thee hnd bd ae preent he embyo i vey elngate exending over abt three-forth of he yo irmferne The gH gr more rapdy a th tage elngaing and trfraing frma inge ba t Pgmentatin beome more obv a melaphre develplng the mid-rna region and t a eer degree alng the atera bdy margint the ta tip Meanophore are at frt abent frm the vertebra regn and mbbd Oipta ad nternaa fontanele are til onp at thi tage Draing f e majr feare f embrygenei re preented n g 12 hrgh19 In eah f the figur the hrizontal line rereent a aJe f ne mm

A the imb bd eongate artilatin ad dgit beme dtnguabe he g beome omehat redd n vera e althgh h dta bran are more dvertiate Dstnt segmentaton of the abdomina regn ocursa the otal fold devop ee appear a eparate ridge overyng the yol awhih oupies the ventra regn between the lmb grdles (Fig. 18). Pig-mentaton rapdly expands ver the head and body regn, csl idges and hind lb; i alos bsn from e forelmbs. Noli,1 gov a 150 dtnct.


17/103

OO O OOO O 2

At leat wo we later e head tes on a defniive hpe, he gafold becomes opious d he fee wel formed Alhough varable, edc-o i gil fiment may e bl porions re grdly reored wia cocomtn fuson of the ostal rdge A tme hort yoIk sal etendsfrom e pospetoral regio. Jut before hatching e yoIk ms is etenallyibe a a light entral extusio Pigmentatio at h tme is weU deveoped wihrecognizabe menophore iridophores d erythrophor The young salamanderoupis the enre cenra cvi of te egg d lies tghtly refed n a edto-il poston (Fg 9)

have o ee be o foow e nre seqee of deveopme reto tme On the bais of comarative ob�evation of h eriod invovedbetween ario nd ovelapping sge in differe ches i sem probableha four o fe monhs are reured for inrao deveopme Th estmate mar to that gven for Ensatia hchozi by EIN 70 Moreover, hemerobat vge pten of the large neay isoecithal yol ad e prolongedntrao period bece a free laal age are consten with obsera-ons of h spei

HATCING

Oe h of 22 eggs nd nothe of 3 were ucesuUy brought troughhathg The hd been maintaned with he atendng adut at o 5 e incoered pas otae partiy fied w moit soil and mo"

Emergnce from he egg was usaUy peceded y spamodic movemensof he yg as it hanged poion wn the cape omet afterwars adde colse exteson of he re ody rupured the cofiingmemrae hch he colased fro lo of itera1 reure suay

the hd or a regos were exosed at fr d he offsrg remaed ill orsee mt efor throwing itse free of he capsule by a seod covutnso Sommes he jtíl opeing 'as arge eough to prmt completemrgece Of 5 es obseed, the perid fro fr upurg o e apsleo emergence vried fro five econds o minue. n one of e es eendays esd bete the hatchg of te frst ad as! eggs Murement ofstandard egth of te hatchg were obaed wt ix hours afe emerge ced raged from 92 to .0 mm aerage 9.9 ± 00 mm)

t ergece th abdome s sll open a1g narrow etral uure

eposg the mal remg yok mass The fragie antoic gill may or mayot or off he ml forc i wy hroug a nrrow opeg of the cpse Their ubequ 15 es1 from brao gi he sol or mos i emrohab or a few hos afer los of he gs lgh r rmn at etera mrg of the guar fold. The firs defte reponse of the ew born io ee coer mo darkened rerea N edency fo aggregtion has beenobered. The tending adt bot females evidenced o inert in he hatchng roes nor n he youg


18/103

3 VSA DE BOGI OC

GROWT

Field eods on gowth were obtined from 9 rpt of 36 maredpeme. OE the a tal oE 87 maI woe apte rods epeent xmonths o 'oe e presenty onsdeed. Iniidual gowth inemets dungperiod oE frm ix mont to ne ye! nged fm zeo to five mm Dung tcfna fed tdy of tober 93 foteen of seen aamnde olleted tPópuation II ad II wee dentifib from toe lips mde p to 6 mothseaHer. Msemets tme E ape ad ast epte fo ese areg n ig Te range of growt demotted by the even mae� in thi

serie ws rom 5 mm (ttind 5 months) to mm (n 17 moths nby even femle fom 3 m by oe ndvdul i 5 ot ad by atheover 6 mnths to 9 mm 5 months

deay, gowh haet hod be ted h bas o epetedmeasements of ndividl of known ges t egular ntevals oe prologedpeiod most popltn, howe" the oppont to omie o esedesiderata s rely avalable Among slamndcs, the m sze o hathngmakes them most mossbe ubjets Eor mrking and thei reave iErequetapte povde mted ormao. Bcse the nos epodutie patten of B. Jbplmt¡ nie poblems e pesente tht h\e not bec eceed in tudes E yli etarop plthodontd No yong of te yer nor Oh-rage ases an be dety sgned Thu, a:l growh incms Cot bappied to ay ge atery wthot is cstblshLg b diret ethods so estmatg g- The oy avaílable predue s to ov the obeved ¡ements to an te Eo ompaon thoghot the now Se nge fo speie My nye and poetion of the nfomaon re thcr bad omen grwth treds at the unvoidable ost of tention t ndvdul varaon

he reltonshp of ompted gowth tes t ne m ze lses s p[ted ig Amg reod obed fo m rgng se f 5 to4

mm wee tht demonsated o measabl growth ove period oE fmsi to ten mn (an addtion spemen dd not nee i s uiginterva of fom ne to fie month These exmpe orcd thogot eyear nd ontribte n expresion oE a patte wi prods oEeggible ze ae i atae wit thoe oE moe ad growh l·thogh ero vlues i a wy vouh fo the obsred wes oE gowth B su·a the ae f 1e e in he deeminton o nnu raes sine 11 nmal oE mre tha te months of reord demnted a ze inese. Ts mates E nnl growth ates mt ede thoe f whih n nrse ws oted.mao aaab fo 75 o he 87 eod how Fg 1 Fo he

males n this goup the aveage nnul growth rate 1S 87 mm ;e3 to 6 for 7 femaes it is 9 3 mm 6 to 7 and o veuvenles 3 58 mm 3 6 Tst fo vane mong thee port a n1 hyothe that no differenes ex mong the hree goups at the99 ofdee ve


19/103

OOY OLOLOSS 3 1

Fg. 2 gs he aeag annual gwh ae f ah n sz las,ming pmns or wc tee wa measuble gro'vth. Alog a magnoí eror i ncaabl, te average wold ve to be td as s!v at oetas f w zro ncrmn ere noted Fg 2 ) Te rod satt-ng ad abe y tnd amg he plocd dicas ha or-lto exiss bewn i a aerge an gowh Wii h ·of 22 ad 62 sandrd ngh fo 75 mls h v: 9 ± 08 w mes om 0.6 to 60 m.

Edene [rm limted 1d obets nt metng m r!o used peiouly sgges oe rd gowh atng five mm p

r durig eid f hacing (vre 1 mm at of mm vrge se: It may a ag dut hs ov mm h:e anual irmt the apah mxim

In pt f eisig imtions, w th ppos o ts o pen a stmate o gt re of B b to establsh so sifor easoablc a o zeag J laosh. Appl t Y K , whee Y = i¡l tdrd engh h adde g t, elped ime ya), and y t snded ngh th d of ,suh a cve i5 imply divd �a appear t b lil etme fo t v

fo bsed pn he ud age gw ) 2 mm p ee. gt Qv cosrcd for B jg 3 eis

a rag-ine ¡rae n an ahmei po DE er" rag e 22nd mm om he hahing � a n mm h am of a of n, or vl of K, is eaed a fiv mm p d shown as curvner i hs prt of h ag. Bod m o iomaoi vailabe wch pems quyg ay mofon pojed ln

SEUA CHRCS

A typial f val speis th Plethooae ad se mmeof te Smndrda B b« se"s lo es he m o lgs bs know lodod nus DeJl1ogitb (BURGER¡ ; HUMPRY43 oweve, te lobe B b ae id o but ned o oas n a xratrpal geS A dege dmnstr o n zand ber f lb fo ach eis in igl nm In 1% o 4 ml o he ste ibi one ob moe han th ote A dife mre ha elobe hs ot be observed. I cs he x uly mall and mar, e o rdg sermatoa

mae male pmaxllay e prrd bond o p marg,and h onpcuy rnaed sn boded by olen rdg


20/103

32 L L

be th tail. NBLE (53) has atrtd crthp tmn a th ro o

hes "hedoc glnd male plehdnds ei uncn n cuhp hben nerrd by RN (54, S) fo to ss Po and by (70) fo E Jhfolz.

In my srs f B bp"la, female ttan mmm stdard lngthof 690 m 40 mm gtr an ht of larges ma e snu is moeded hn n m d soab gov ar jncnsicuo Mmbnosd h mag of th cac. Nr-mnl grvd fm cme lags size clss of al 'pecmens ose

Swollen coca ps ae charteitc o b sex whn n brngcndtion Ths manfesaon hweve ndive only f e cul eed·g pid f he indidua ad abnce cnnot be sed as gn critronfr rrdcte mmtrty t ds sr s usefl chrct i dmnngh duton of the brdng cy n th ppton. hoghot th pd od tY ms nd ema wre obsrd n this cnto

POPO DYS

Afte plmnry nvstgan to detmn he tnt of otraty B. bpl sudies on ivi yes wer iid ne hu fo dak ndcnnu un a srie A eas tw sts dffrent vton wer vsted ee tw hur during the ngh. A ota f 3 ste sts we md Tinsige he psile nfuence f soal chnges upon cty, hse tdewr conducd Otobe Decemer Mah, My ad Jy 961 and 9.

B sba s coplely nocnal. Diunal civiy hs n induced osimted to dgree by rfaU m eces f ethdontds uh Deog-1h oe 1o!e HOJ 4, bt recptat h not

bseed ifuene ativy peiods o samndes n h ro bre honst of daes. Deep wthn e fot, or n m densy etated qbrds i is psle h lghtng is s educed eve dung dight hs se durna acvty my b smad by ncrsed amohrc mour On can Mch 1 962, an exposed uvele ws colected n tlus slopena the mmt btween 2 30 nd :00 hre hd ben no rf dunth day. A the me of catre he gd rfce tmperatre regster 2.C; oud cve limied visiy 20 m d he lave hmdi ws 00%.It hod be noted, hweve hat suh condn are ypc n Cerro py dung lae monng y afeno.

Ativt riods may etend hughot !e nght egng st trdus and temning wih he fs dcenl lgh of dawn. Vbl mgnce imes we evident wtín he sme sm aa dfn nht dt frn sample a on the sme nght e o inl mergce rgedfom 6 PM o :30 PM GMT6). A ths laitde N he s only


21/103


22/103

DE BOI OC

aratios. he grad rese of acviy of the smders to a sige m

mm pk was modified by he onset of preipito, a whch me h otanmber f exposed nmls bme reded Uon sai of rfa eewas y a eond pa whih may or may ot have ben greer h the fírst,dependg uon im of rence ntesiy and dration of rain.

The hrtertcs of aiy patter cn be exmned n Figres 24 nd25 whh nde iforaton obtned a he ts of Popation 11 29 m)o the night f Stember 18·19, 1961 ad from Poaio 11 ner e smmon May 12-13 1962 oiar nterfae temperas ky codons, retvehuidies and ercntages of ota acivy re gven for eah wo hr nterval

rom beor dk utl afer dawn.In e mod uve (g 24 ctivy wa fír obsered at 7 :0 Mand he sge mmm k ple bween 30 and 0:00 PM, althogh soemades were acve at he last re_daw isit at 4 00 AM. N ran r-red dring he ight b grod leve clouds mnaid a tured atm-sphre un abo 800 PM after whh the kis rmained ear

n comprio, the ifene of precpton o degree oE city diaed n Fg 25 The frs ms emerged jst at dar and the frspa curd a 8:30 rom tht tme il 1230 Al the leve! ws redcedbt agai reased to a seond lsse pea at 2 30 AM. Here he nii acty

pea reahed s maximm at a relte hmdy of 80%. The ivi ee wreduced s hmdy deeaed Afer the declie of atiy a sod pk red immdtey foowng erod of ght r and may have been simu·lted by it A hgher leve! mght have bee aned oowing he 2 30 M o·ation bu dug the 430 M vsit he early ght o daw oly reded atiy

BeCa1e te illnaion ad hadig ec"ary o detify smnswold hav flueed ther ay a o fble to deeme te auallengh of me most salmders remn ave dung a sige gh. eain

ndvidals, however were aly cogzed by sie n color.Noe were atve for more h wo sdy ievals or a perd of ou horsduig e ight f he me betw sdy itevals is added to the fírst Orat ghtg t wold idene a mamm aviy perio� of s hos

Two predres wre folowed to sess the ata prprtion of faceaviy poplaons 0 te nght of May 12-13 16, a smpe was sdiedi e viiy of opation A ad at e interva hroghot the ght allacve amals were coleted A snrse, afer comete cesion f vyhe ara was horoghy eamied and all avaiabe aamders aen f onlythi rfae or avae popuaion codered 117 7 of the tol 'peof 60 ndivdl s acive

Dung activy sud at th mred popato sites o anmas ereoleced erefore h terva dd soe amals that were obered ad recrded more tha once. To avod ay mltip reords of ai idda have ed the perid of gratt atviy dng ay vit o esma te hedegre of acvy i he populaton he r of actve dvdua omped


23/103

ECOLY OF BOlTOG10SSA UBPMAA COTA RI

wth the total oberved alamaders a mared popato wthin a 'e of thgh d gve actity eves fom 98 to 24.6% f obseed avty evelre ompared wth predicted umbers of ndvida n the ty areas ompedro capturereaptue data fo moth dug which atviy �udies were madehe propoton of atvity i rdued to a rage of 000 to 000% of the totaloplaton

At he maxmm t a qartr of th popuation atve in ayone period No orreaton the teniy oE avy with saonal mat re-gme ha be demotrated although lol weathe ondtion do infene

emegnce On any gve ght. Fthmoe,1

hav obtaned o evdce whhcoreats dffrece perodit o evels o atvty wth ex or siz oí nd-vdual.

MOVENTS

Throughout te rage o B mbpata on the Cero de a M1erte thees a hgh caryig capay utaned by the avaiabe hta. Durna retea areprataly aways availabe to the anmal wtn a dtace that can be travereddug a noal acvty prod Spaia utizaton of the to habtat thre

dmeional Potetial retreat are ot ony mmedjatey beah srfac coebut al nude ubtean eel to dpth of probably a meer o more in aludepots uring peod of avty smander prmarly utize the godsurfae bt ao have bee obered mg o rock d log or in hry arborea habt o te pee ha aready ben mentond

Colleion of subsurfac spemen offer oe information a to thedegree of vetica movemen wth te ubtratum. Qe cvaton of a quaremter ar proded negative r/t, whil anothr proded 25 divdal tkefro cave ad revces ro so to a depth of 30 n o ocasofo:teen adut wee tak from withn a rog g appromately 2 met egh ad diameter

1 ume tht threedmensioa movmt ar taing pace dg ormaaviy o !h see wthi t home rage Reeted catue of e ame di-vdul at the ame te after intrvals n whch t as now to b abent an oy acounted ír by ubterrea ativity

Meareet of movemes have bee based upon staght distancew refuga Ahough ot takg it account th t(dimenioal areof actvty o wandrg betwen rteats, tese recod or a bas o evaatio o the ompative degree of mobility oí idivida. Reords wer obtained

from a tota oí 269 mo of 194 maed anmal preet at the ve atalpopation ts The did re eaptred from oe to a many a eighttm

By assgnig th mos of B to drete ieraI n time anddstce, tatstiaJ analyss of th reatioship of tee wo factors a e made.Mesured dstanes separate move rathr ta total dtace overd o etdace from a gve wee ploed on mer u agant h aal


24/103

tTA DE BÓLOGIA ROP

TABLE

Movements B. ubpamat expreed n mohy nlevas. The n hyphes sueIha dJance moed ae ando wh ec o im Chqu ae aepe ¡h 9% Odee ee a N 2 dg"es 0 fdom Aek * in/es ejo

MtlyIn'aJ

1Z36

i m�c

287.13.05

021

0 he hpohs

Fr.

1231460

'rABL

PossoNW!bs

'

87

19

5

7.G1

6964 •69 82

' "

.2 88 ·

Comp(1ae eage 0 dsaes ed by es, e nd venes. The hyphsi aSSms gnan deee ee h hee grops Cha

es e epd h 9% odene ee one dege o eedo

Maes Fem:e vns Chi-squr

5.4; 3 6

2l .1. ' 566

AL

ompaa ege 0 me of ;OS o aes, ees d en. Th h)pohei !e 0 gnan dfeen en an he hee gop Ch-:

a ae aeped a he onfdne e oe dege 0 eedo

Ma

.1

mas

U

u

1

éhsar

0OZ000007


25/103

ECGY B IN O r 37

pod o tme monhs betwen o captures OE otl mos obseed228 w over peods fom oe to sx monts wee ompd sepaey th Posso dsrbuo ob maio of nre o ndomnss movemen. able 3 icudes petinet aus dd fo te alcuon

Te vlue of s he vrg de meers ed fo anmlsdung tat time eal suae comtatios ae deved by u oE squaddeaons fom te Poson mbe fo ea dane e n-vas te distanes movd ejed beig random on te bs oE Chsquras N deges oE Edom Eo h saes ea gety eeeding te95 % onEidene el. f th dstance of momnts inase rndmly wt

te would dce a ndidas do not o}' an deEied Thedsaes re rndo owe plyg a hm ge s id.ed cods o moet of unsd jeis « 30 mm sndard

Jeg) udd 15 anmas. OE o oE 17 mos emes ere 010nd m I ad males he imits oE 75 obed moes of 6 spmesraged fom 0.10 o 503 m and for moes oE 16 Emales bte 005ad 35.13 To aa infomaon dtances ve be ssgned nts of e mete t fqences of ocuene fo jvnils mles ad fmas(g. 26) s oedu dsegad ay m nl.

Amog veles te avege ds oE aU moves s 285 ± 084 m

w vege of 13 mov per ddul Adul males verage dsnof 543 0.97 m wth 9 moes pe T magde of ea movey fas verg omew Jess 3.71 + 042 m tan for mles but h vr-ge umr of moves pe ndvdl s ligty greate 153.

Averags of he distne of moemen t nu ypotess bng te diffeene magnds no sgnfin beee uvnles mal adfemaes are treated n Te 4 ea oE these ompaisons te l hypo-hess s subtantted t e 95% evel Eo ne dgee o fredom

On the ssuo at no signan xsts n diffeen of the m_ of movs per dd uvnles, fme d males Chsque evu-o ¡gai substntts t u hypothess a he 95% obbiy lee wh onedegree o frdom se be 5 )

elatiosp of dsans moed o sizes of saJmandes an owbe xamied n Fg. 27 he ge of obseed dstanes oE moes for eacto se ls s ploted for uenes males and fmaes W ten s els, mles of 30 to 39 sandd lengh ae epeseed by 2 eords; ose betwee 0 d 49 by 57 fom 50 o 59 by fe dhose ov 60 by oe Fmals of 30 o 9 iluded 4 obsevatinshose from 40 o 49 ilude 60; fom 50 o 59 6; nd abov 60 13. Interals Eo whih tee wee fe cods de

mostated be higher ave-gs. The pe of ese rsuls doc> t sgge any orlo bewen siz ofh anmal nd aveage mgniude of movment

he exent of mobliy size lses of B balm s peseted Fg.28 Jottd a o ineas oE andrd gh, agi fo groups, areh vera� me oE recorded move. Hre so te ves eneay el


26/103

8 EVSTA DE JOGI iCL

sze d of animal mo freqeny cape. T inconte ovrappng e poxty oE vae no evece ay coelao beween i ae amobliy

HOME RAGE

Th lte nfomaio avaab o aamar hoe · un Tape of amphibian ecoogy ha ben coee ony incnay by mo work who hv ariabe eam To ef h concep have epenheaviy pon h ierpetao o BURT (3) , 0 an FITH 25

Cri pe oE moeme hav be conee o be excve of hom

age ar UT 1 o exampe ha a that "occaoa ae o !h aa perhap ploaor ae hol not be coni a paof h home age. In hi wok o Ra oa in Yeowo Pak UNE74 t yic migaio an ap popaton h a pa of e avity ag b byo e omal mi of wha he efee o a a iho age> Sch eaoa wanng ae w kown in aH erreal e-bae gop b cognze ha iinco beween rap h empoa exion are araterze more in magnt o ime rath hnae (omng o coe ongag mgaon CUNIGAM (7) hpointe o hat he enia eenc beween emporary exee ore ah poio o w home age i hat of tme

A conceve he home age of B sbpa de ha aawh whch te va ptety cae it orma roe he com-mni. no being mgaion o aggregaon o n thi pec he oyati whh j no con o be whin a hoe ange i mo ha maybe aociae wih peron of h popao. n he abece o ay knowneroa, an conieig ta h i elom a lmaon on hbita pace a tac betwen o poece cover in xce he averag anf oe he ho age vew a beng eena1y [ar.

1 hae aop in pa he meho or eteno of home rangea fin by (25) in whch two erent yp f mearmt ca : obe anc bew wo ccve ite of pre ofa iial, an 2 the loge (ne) ace bween ay wo captre ponT proc off h avaage of perming termnation o the avage hom ange o mbe f he popao m mte il capa a compaon o he se oE mnmm ho ang aa E iv imat om he n o oa oe ance. ran iaatage a inheren any appcaio o !h mem They o no oer any incaion oth x o whh oe pa o hom ag ae bect o geae hanohe an he ragh i bew�en wo pot o cape repeenly he mimm Ol ne chage in poiton

e aage home ag ae i compe a he aa o ccl ( ')ng gh line ance (ai ecoe in 290 recaptre o whih mab oion hae > 005 m) we avaiabe. Rai exrem 005


27/103

[ ECOtOGY LOLOSSA PALMAA CA 39

0.39 m) ae heeore eqal o ose obaned or sepae mves; aea eremes

woud be' fm 0008 to 797.00 m. e aveage hoe ange adus s 37 1.19 m and e avage aea i 436 4. s m As no sasca sgncandeenes were ound n omprng moves o juvnes, maes nd emaehee aveg sho unom app

In he absne o n onador evdence 1 am pg e genehaace o home anges o exaropc pehodonds o be ssen1 he smeas a o B. bp¿m. Masremens om he olowng ans have beenonvered mees In ESi (EBBN 69) sgncan deenes wee ob-seved n e aveage home ange ad o adu mes (95 m and emJes

(10.1 m) gvng aveage home ange aeas o 1,96 ad 3 s m e·speve $ deemnaon has o be easbe n sdes on movmens oBcseps aena o B. paúc ENICKSO (3 gves h veage"rsng dis o he or sees a 1 5 whh wod be an aveagehoe age aea o 71 s m

Anmas epd moe han once a he sme se e no e measureo he een o home range hogh hs knd o noman s vuabe nndang he degee o uzaon o hoe se The seden ne oípehodond smes has ben commeed upon n eah o he boye papes b s somewh dc o eae em n he wo repos menoned

boe vagi was msed wh eeence o hanges o cver wou egdo sz o he suae obecs an movemens oung neh hem BBNS(9 observed 39 o 5 maked Es;a ha emned nde he same coveom one o ve monhs dng our eas o sd In hs wok on Brhp, NCKSO (35 odued eave age ve board whhpemed onsdeable poson hanges beneh hem I s possbe hs vaue59% o 133 maked anmas whh dd no change cove n 24 monhs ma havebe nfluenced b s acor and wod heee nsevave O 375 epues hp pa NNlGHAM eps h 9 move

o ohe oveEvdene o e sedena naure o Bigosa pam s bsed upon22 o 406 5) ecapes n whh e anms had no hanged posmoe han 0.05 m who egadng e se o ve n peods up o enmons. No sgnan deen cod be noed as o se or se. An ademae aen ne mes dung a weve mon ped a hanged aon bewee eah obsrvaon bu e s o al moves aled onl 6 mUsa he haace o movemen dd no demonsae h degee o cvwh s sho oa dsane howeve he grees dsane mved b andvdua o wh hee wee wo o moe ecapes was sg e n

21 mOM BEHAVIO

Fed epemens were conduced on nne mked adus ( emaesand ee maes o deermne he naue and degree o homng nsne salmanders were pad a deen diens nd dsans wQ


28/103

40

to elv m írm he points of origna capure, and hes s wr hkda six wky intevs oowing diacemen. In only oe innc w a d-

placd a oud in is oginal si, frm whic t hd b moved trm Wk befor. Th fe wa agn r�movd nd acd evn m wyn he am ireion bu nve! wa observd rrn. Aough no ohr -placed BJ rernd vera had bn acd nrer o e cature po.

es ew rcord· can n mor n ggs h hmng bhavor pooly dvloped n B. sbpalm Howv, t ack of fdy egg cuthatendanc s we a h se wih whi adu bandon thm, ofs suppor¡ng nferetl videnc

GGGN

eow vel o 2500 m population dns ar ow and do rr r more alamandr ound in cse poimiy wihn e sme rug.n midrangs nd upwd to he sut ara as dniies nreae w Oavorabe envona conion h ourence o agggaons becoms morefrquen Above 3000 m he ndg of o fore animas under rokno more tn 5 cm n deter COon

No ndicaion of sua behvior ha bn obvd n th compoono aggrgaion. Th group wre composd o l mes a fales or vayngnubrs of boh es wih equal rquency By ollowng he momnt omar ndvdus te compon of ggregaos a oberved to changemrdly n ony hre o 90 03% csons wre the ame wo mastogeher more han onc n me refugum

Durng he dry eaSOn on h erro, h ground ufc bcomes cedand almander are oud ly undr deep etd debr nd ros wher mo_tr has bn etand. Aggrgaos r inensid during his For mpebeeen eptembr, 96 d bruy, 92 ony 7 pemens hd ben oundt sit of Popuaion n Marh, howvr, 5 n ndvidual wre mard, o whih were in lolzed ggrgaon By epmta mth hv odht ths bhvo s of soe urvv vntg in oweng es of wr 10s nndivduls by reducng h eposed body uce durng ses condio

At es of oc indation a imi yp of behavior s eid As s-mrgenc tolerances r died n nohe secon o thi paper it wll ufcto stt tht avodnc of xcesve ost wl ocsony force sal djds h sm ee

DN

o mas of phbn popon dnt hv ben dvd fmcpure_rptu daa ulizg so orm o h proporton1 ind md.For prinent traent and evauaon of such procedures h ar reerredto th sries o pap by TURER (74, e¡ Y.) Available normaion on dns of smander popultons i notbl ony for he lmed number of sd"tat del wth ths sp of poputo cology d arty o thnu<ply


29/103

RI 1

B bfalt utilize i habita to a wide degee n iew of this fa

he owes (or rde) desities ad eoogic dies ae eqaed Eideefor etation of these pametes a been obtained fom a total of 290 eap·es of 936 maked slamanders at fie popation sites ove a one yea peiodObervatios of relatie densi ae of moe lmited vale, but hey ca be use-fI i detemning geeal ditrbuton paers Rlaive density esaes arebased upon 69 specmes tken from 60 quaey quadat sampls in the sdyaea

here is an amos cotat pate of decig dsity wih ower evaion he ale fo ach quadat from wch specmens wee taken is given

in able 6 and is obtained by compung elaie desity as he nber of indiidas pe sq. m. No aimas wee ever coIeed i 20 quad sampes fromeleatons below 2438 m on he nohwe pe r 2591 r on the othernope n ese qadats, densities ae expessed a < 000 whic epeene propoon of one q m to the toa quadat sie or /186 q m and hallows for he posibility of B. subpmta ocuig in he geeal aea a veyw desities he extemes of relative densii in aH quadats whee the specieshas been oIected ge fom 0003 o 03333 e sq m. The aveages of fousampes fom each eeatio are beween < 00282 o < 0.181 idivdas perq m Variation eisg corelation o alitde with denity ae ot sigificant Alhough attempts were made to equae he physica ondiions of adras ea sampg e csi of ing dffere ses ndoedy infenced relts Aso a we r more was quied o complee each seqence oísamples ad local changes in weahe onions are possibly efeed i hda spite o hes souces of eo, he coeatio conspios

Beter pespecie of the altitdina distibuo of B I1bpmat caD begaied when the rlaie densities of animas oeted fom quadas ae plotd against biotemeatue ee on he Cero de a uete Fig 29 inudes qadra apes The gret desiies occ in he ower bioempea

ónge. cases whee here wee 01 or more aimals pe sq m bieperature were below 06° C. f 6 spes fom sites above 5' C oy 9 %

were eve ocupied by B. bpimat drig e ye ad e high desirepreseted was 0075 per q m; eql o the owes denit eve eve ecordeda he sit

Salamander popaons on he ero de la Mee are subjet o onspiuo hanges in densiies The impa of e JaayMah dry eason is byfa e most intense ad lowest face desies oc duig is time Wihthe on of Api ais (efer to Figs. and he anmals move o the suface fom heir subterraea eugia Fcaos ao oc in reponse o shor-

e and less iese dry perod as the Veanillo de an an in ly ad Agste mo skg eamle changes in rlatie nbers can be se ear thes where optma habita codions ad geaes densies peva sampes taken from a series of quatery quadats a 3048 m eation o he sohsope, bes o indiiduas coleed raged from o 62 represeig sufaedensiies of from < 0005 o 03 mas e q m easonl variain


30/103

42 REVS D BlOGA l

ALE

Rv d 01 f B subpamata

(dmd ¡m q V :

wzb dZ . m. [Alz * e n e 1 m ]

!P

NNNNN

mS

ohyAg

S

Pd

BnU

dgbJ

nn&

BgJb

Evaoni Mts M:ch J D Ae

438 4 29 3


31/103


32/103

4 RVSTA DE BIOLA TOPC

Popato I ad dstes raged frm 7 66 nd 2 t

6 res:ciy e asee o correatn n time Qrence eains odu oa climatic neen iew o he naue o the deind sy as avaHa data 01 deit¡eof extratrpcal pethdtds apper t be cont oer-etimas Fr ppe coaro, de deiti of B. subpma c be mpued 0 e asiof the actual sdy ar rher a a adjtd vaue d by dditi oh oe re radis_ Uoced dtes o fm 77 sander pr s m repree te etremes of oserved anual aerag otaed drigmy sdies Table 7 sarze e parate daa aae fo for ospc ad euats e nmers f didals sq m NDERSON (2)

udied ppuats o ds gubi and Baho uu eiands o S rasc Bay" Caforna The aerage dy of our tmtese obtaed r Ai wa 892 per re s remarkabe emae o 663Bhoss per acre admedly 'ecessive whch e arited a loclpplaio shit; however, e nideed an emat of 86 de peare b reliabe.

the obrved avy level f B subpma/ are in any way dtveof ow eergy dad y ee mtne ecerm, te e factors lige mer of dvda wold appar to e deniy-ndependent; mos prbaly, paal rquremet and avaabe micros pes wii the ditb

raneAGE CLASS

Age a ara of amphiian popuatn have uuy een aedp data from arg amp oeed wth a ho perid d e ie asesmpared as n BNIKOv" di On Rao ibius ad Bombiombi (6) Sverl peotid amaders NSON ;GR 28 29 OE 9 E and E 6 6 have ided an nalysf e omion o popaios, a in reerne t e propon o yongad and adts. nvaraby yng anima ae seprabe o e ba f

ie. s the age asses ncreaed e mer f mae aduts exceeded te younger grop ad oeaed te rg f te istogr Tese merggpaers adt ave mpicted e ntepetato ieage as eatihips of oder indivdals Mesurements B. ubpa ppuatin ampeproide o dire edence rm whh age emae can e made s hwfrom he data n g 31 weeer exte�ie and yea! rd fmat saalabe a paer sie frqees i dmonrated t acks cositet ndao of age grpins

estiar have een ued y HUMPREY (43) nd RC (6o tabs age as in Dmghus. y exmation of istlgica materiaf Dmog7/s f HUMRY 3 determed a males w omre esticar Joes were n heir third ean eua activy d hat eaeuet le reired w year to deveop fy. Using he sm me nterv A 6 defied a of mohs av e ob


33/103

ECGY JtTGLSSA SPALMA I STA 45

eee hre n fur years; wo obes represenng fiv o i years ad Ihrel indicative of sven eigh years. n Ihe five es inuded' in his stdy ales ared y e fourh yr afer y had een depsied as eggs.

Fig 32 nlud a sample of 140 B Jbpmal aes aen duringJue of 962 Te wo m inera ws chosen wi he purose of avoidingany bas i favor of he proosed growh crve (ig 23) Singeobed maesin this spe demonraed a oda grup a he idpoi n doube·obed les the mode ccrred a 47 m, wi a second pak at 55 wiee ode of Ihe Ireeobed speciens ws at 55 m Thes mods may b coicieta afats, b th obviouly confor o the esmaed 2. m per yargrwh increens deried from arked ad raptred anms ad coresodlms exacly o inteals at e nnth leventh and thirnh yers, wihoutccounig for any rnge óE vaiaio f he biennial paen of apearace fteiar obe correcly aplies o is coinuousy erdung roical eesen te aeag represned by e growh rve are indirey sbaiated

SUAL

The caacity for rerodution is chraerized in a by h reeneof aur serm in he testes and Wffian dus d in fmaes by h a-prnce of heed conolued vdus d lage a > 25 da). bh es miy is atad oer a farly wide ange of ag and sie.

From mcope ides of gonad issue or 54 B bpafmat aesbe 2 and 65 mm saad g he rina has of sertogesispree are reogizabe. A snge me of 26 m represens he es idivid-ul attainng auriy Above his se he propotion f are aes generalyicreases to e 42 mm size ca�s aoe whch a specimens eed were pdUg sperm A size direce of 6 mm eis :e e ont ad omeionof sexal mriy in he mae eeent of he aon

The ocrrence í maur s i he va deres apes o be sim-eous with thir first oence in he hough n % o e isuepraraions erata ser coud t b ound in aure aimas issible ha is is due o artifa rsing from the monng ehques ·ploed. f a ino fation ds r n he reprduve actiiy of alesr a period o prm enion xiss, obsertons d n indicate ny craio wi eier size or sean

ORGN (56) used icened conv!uted ovidus e crieron of feale auriy bu and (60 onsidered hes scurs to sutje lO rere resrpn, whih tme ey appeared straght and narrowbues aeaig wih Ie perios of breedig aiviy n a sme of 35 Bbp females (oted a 2 m intevas i ig 33) he sie cassbeween 38 ad 54 m inuded 58 individuas wi and wihout ature ovi·dus Faes boe 38 mm deonstrated a generaly conssen increase in hprortion of h sample t pessed is charaer. Of 16 feaes n e 540to 559 nteval 5 % ad ever ¡id eggs nor did te condiio of


34/103

6 IT D BQA RA

ovids indcate tat they were in a physiologicl condion to do so A 66

pemens of 56 mm o more were mature by ths cteion, athough mos oChe te ov cold be ssed s mnue ( < 0 dia) and dd no demon-ste ay yok acultion. Fve of the !r grop (7.6% wre obvslyspe s certain hat maure ovidu pesist once they have develope.

Th condiion o he ovidus does no dequaely estabish sexu ma-turiy in femles onsideron mst lso be given to si ad age o the nwhen ovpton f occus. Althug mate oviduc were obsvd in fe·maes of 38 mm he ma ndivduals noted gravid or broodng were inhe 42439 size ass Among 22 speims exmned in he .0 o 9

se clss 2 had mate ovidu; tee were obvouy grvd and ne wasbrooding eggs. he minimum growt increme of individa beween heonset o mauriy ( e., with matre oviduts) and he perod o egg laying ws6 mm Gavid or brdng fmaes were reprend n al ie lsss above 4mm and a11 emales o 5 o large showd eidence o hving peviodeposited ggs o were n a onío to do so hereor sexl maturiy ifmas ocrred soe ime wihin a growt inerva o 2

he rregr pattern of te peentage of ¡neae beten sz extremes wh mariy ocred ws pobaby a rest of the lmid smpe at easize inevl and variab physologica condiion of indvidas No singlemoda size s was pren in eiher mes or f


35/103

COOY OP OUOLOA LMAA COA

years. Data ofr in th arr ratmt o xua' ubtatiaé hi

tima a gg hat a wo ar r i probabl th minmm rqrdbwn cciv vi gg cuth.

Th vaati i ag at maturt bh x B sbpalá mranab in w th tma 8 ar ta an Rtar matritin ma ma b a rc h tm rquir r t rav hgh nrgma m b abrain o vica c n b a rqr or trago nrin in th va rg rma t ik, t Iha h truct matur ma currng a a m wh aU ma uat ag armatr wu nhac th robabit xua ncut an rructvcc.

SEX RATIO

Evauan o th comin B t oatn i ba ic rm quartr qarat O 695 >cmn 68 cu b xa trmin b ct Th x rat at a am ro t n Tab8 tia rrn a o t ratnh a Ch-ar vau btatath hth at th 5% coinc v Ír o gr o rom

TABLE

Chi-qe e 1 Jhe ro 0 m lo fede n popo o sbpalmt 4eee b ¡he te/y . The n hypotheú a one-tooe ex o' Tbe

wce ete a Ihe 9 % �ofene ev! oe eee em

Monh N FemaJe

rc 39 5n 14 2ember Dmb

TOTAL 336 35

o

9726179

1

o

025303

.4933

CiSqur

8.3

Svra i o thti aamar ha bn rrt b r nvigao o hw caU ignica irc rm a balacx rato Th rnc hav bn brv b n avr o h ma a Aneide¡ enem (D, 8) th a in Pe/hodn vhicuu, Pd1ni (D UMA 21 an P ynee ( 59); r hagig aaU inehdn inreu T a BINGHAM 72. n h ti t gnu Deghus G (5 un Ihr th incu v c o mtratgncant rc h avr h ma

Numro acor hav bn cir a orati a gr rutig n iait i th xua comiton ht at. Brig m o o c ar mr i uctb t ratn (Dg-thush gra ma ma b mr crtiv hn a amg rrr i itr·


36/103

48 �VSA DE ·BLO OP,e

ded (Anies); n Plhd cnr sexl eene n esol avta'e aa,enty sgnifn; the auat hbiat oied by the femae n sospeis of Dgahu durng brooding i a more sere envronmn hn t�terret,l one o he on·b,odng femae; ak of eeding on te part o an attendig male esung in grater useptiity o he igo of oveneringmay be a sexally elee fao

None of hese aues a to ay signiia degree in popuaios of Bubat Femae ith egg uche dmonstra a sape repose no markedy dffeet from other indvd. Mohabits uiized o etig oferoptma habitat onditio nd ae ommony freuened by noreproducing

membes of te poplaton Sdie 0 atvty ye and movemets hve ofered no evidene o indiae hat die o s


37/103

op BOLOS UMA COA l 49

unl doton. A u pod of wo year i ud b t dffena!wn maron o b ovd and yog anma o fond d-g Fg. 33) Pobabl laer priod i ore aura, fo h was alo mnmum tmd to b ud for the enrgen of gg. Thi na a ypa adu maJ a of dosg vy o ehng aury a hc ag of n Gvn h tmatd axmu urv agof gn ya b m pona od be u a avagng225 gg o offng

An mat of erodv pona! a ouo ve! may ontd om feld daa Of 345 fa coHd n quada 3 wgravd ad en r boodng If on he n (2.9 %) of th oa falea ondd a podv ompmn of b popuaon b would fia low Sne no xavao wa don durng i lngo of t o lkly ntng ained t. Addton of 3 grvdfma n 93 % ) od robbl ke ma o g a gg may b ed a la gh onh A mor eonable ba f d-mnng odv oal oad y omng pn fm kn he m gt o 3 % wt b broodng fma and gve a oal of5 % Th dur dd ncoore gra arn of rr b

ano aoddO h ampon a h 18 broodng and en fema r odug a oa of ofpng nl of femals X avg z o gg uhn fro a o of ae ad fal tb a 58.3 % 05/65of ouao ig addd arl T va een madanua paa pon rod cc of b popaon

SURVIOH A MOlY

h ' mot mpfed ehd o ang motal at dr-

mn popon o mad ama p on yea aft ca Oí 3ndvida a o�ao I on th r found ft na If all oftb ogal mpl w aly a k h oty Old be 0 ear = 00 Aogh f rod fom o ouao r moled n nbr ro a ra obnd

To cd b f amnaon o aa B. ubpl' veal rie mt f acd: ( h mad aga !aonhp a vd; ( eodu ad dh a a qual; 3 nodffn n oai ar ibed b he e ad no nbaadgraion rn

ul rduco

oduon

DEh

populatonORGAN (5) atmpd b dmnaon of vop a ni ie oE DeSog1¡th izng h chara oE iobd ee annolang h ag a ro HUMPEY'S (43) te 0 th ba oíona rodcve pattn D fSCI! h later wor wa al o agan aolu nal o tw ya wn h ucv fomaton ca Io� a o may


38/103

Te onfali aionshps of ega a Bo/ügSJa mke

e application of Og's eures to e prese stuy an obvous oieaion. Howeve, beause of the acyic reproduve patte ad appar aiely sow gow ates in B. Jbpama the reslts of ay oprisons suivorshi with gnah on the basis oE tticula obig mus be cautiousy regarded. coosite o qua cords has shown a wie ange ofoveap the sies a hi a gi\e nbe o suar bs ay Dcur ostic the extrs of size diffeen at which le may fo nw bsdurg he a, he ue smp pesned i Fg 3 ws ta as mos apiable olowg e assption tht a!l woloed mal h>ve psse hroughe one-lobe sage d a hee_lobed maes through the olobe tge, he

eaiv s of ese group ay ffe ei fo ipretato of specfag survivorshi aes. To eque t rs of cpur of t gsonly adu anials ae inuded he 0 % aturation eel stabihe re"iousy iuds as duls a mas i th m 3 ipoit) ad p.

f as n the o-oe cas, srve o cqure to obs, agive iei survi! rat 042 bet he ith nd eeveh yeas (asestia Eom e mods) o a anua rate qual to e squre rot o hva!) of 0. f the si eeIobe secims pren a ue poortio oEsiors o h twoIobed gou hen the biial sivoship rt o 00

s an annual at of 09 fo the eevh hroug tieeth years u bemhasied, hower hat hese estima es offer only a genral c fSUviVShip evls h total age angs of the thee goups aot be lueEuhe.

Ee tabes an svivosi cus e diicul to esablish o anmsof conieable longeviy us e nb of deat is a Eiy chaaistag ih eh age gup Becaus of �e absee of ay eas wh perma easonaby accrae etimaion of he orion of maue anias in thpoptio a the sggesed advaed age a whc atuity curs i is oaia o aep thei onstrcon

A moe elaboae saisica ethod or deeinaion oE h focapureecaptr data h b develope by L ad CH (0) BeauseoE e nuerous euie assumions riosly ste A an LE 19)uestio e advisabity oE alyig coplcaed aeaica ceures o nat-

ua populations s one of the afoeentio oniions ae in confi wihthe evinc n B bpaaa how, i ses isabe o atep he aicao oE is meho.

1 have uiized e theepoi ethod oE CHIlY 0)whic emis an esmation of e rae Eom inEotion a three ple

erios oE eqa itral i is a ou moths) and aows for eniyfutuations wiho bcoing ahemtiy wily. norporatig te dataom arked ouJao suvivorsh (P is ress as P 01 2+1 02:

Rohe euas he ube oE aked aJaanes eaure a he seco


39/103

DOlLO UA1' O

spling perod; m02 Ihe med imals from te frst sple not reprdntl e thrd; "2 anmls marked n Ih ft smple nd rcpred "at thseond nd third pu those mred t Ie second smpl nd recapted th third; ' qus the total mrked specmens tken n the hd smp!e ul,he nmrked anma ten in h seond spl and R equas Ihe ttl nbr of e' fi[t smle Sbsttíng vles, P 8 5 + 28 2 02

6 (7)p ye nd is tred s contnt for ll ge grous.

CAUSES OP MORTAL

ere seems to be lile predon upon B bpala n e Ver�ons brd nd smll mmmls m ke soe toll, although no drctevdnc is úlbl Of mphibn R pie nd Eleherodacylus spsonly r ponds nd der rks below 2800 m e known vo-ious and dsmne ding hbs nd and leptodclid fogs sggestLe possbli of thse spces s predos h mJl gund snake, N"ia ox- hs bn cpred in hbtats n the Cero tht oud indice possiblepredto reltonshp wit B. 1bpaa, alogh e wo species hae not befond togeher

roughout te vecl ange of i dstibuion on he Cerro bpm is found wh wo seies of lzads. mnt/, an nguid (m-mny hares he sme terrestrl microhbt wh ts slmader, ·as doesSceop mdhc. The tt is prmly nsecvorous and s doubful s predtor, alhogh jvenl ma cmpte for fod tems th B sbpmaA cpte Br 01c t an dult B bmt whn 4 hous ter thelrds pe

is possbl tt se durnl bid spece sh a Tf ngeceJIe blak robn inude samnders in her de The omlementa nure of

ctt yles wold pem only mited epusulr pedation Hower, soeslaaders be found n th sufae a magns of roks and 10gs duingte dy Te peved in Elsnti by SEBBN 70

O one occson n ttndng femle and e mss disappr fm enestng sie cse of vious obsrvatons the abence of he female ws nossng but h ompee absenc o ny remnans of uch whi hadben recenl) deposted could only b ounted for b ts trnfer to anoherite or b compte desrton

Pates obsred lO fest B sbpml inlude nmatoe woms foundn the dgestve trt nd mseneries of 03 % ( of 70) of th specmensexined. Acnthoephln le occred n the bdomina cvi of % of


40/103

I DE Ib 'OL

his pe No etopaes wer ober�d d no micrcopic sdies w

made of the tuesThe khood of feng o desccton do no pp a eioumortty fcto Tempera below feing e rre and ocazd j occurencon he Ceo. Avodnce of ee condons cn y be cmped by teat below the ufce. At owr eevtions deiccaton may b a rlativly moeerious theat ding cti times c ye although subuac etrea serv n efcti scap from th advncing d eson . 0 Q sio May 14 192 dead dut wa found under ock a the summit t s not poib o epanhe reason fo det nO to dtrmine if t observed dsiccatd oto wthe caus or rt of death

The imact of human ay pon ppulton of B. bpd is di to sse on ong rnge pespve. Lumberg d othe prcce whicnvove rmov oE much c the ors wl b deleteos, epclly whn it ccompnied by the of nd fo frmng ad gng Howeve the adaptobilt shown by B 1bpa utizing mnmade habtats conpiuousRoad cts fis nd tals sloe rsulting from roadbuidin activts are dnsinhabited epeclly in higher eletions Debri, such a lumb nd buidnmate dtrioating wher t h abandod at conton camps oí moen en yrs go o been tied by ese mander. I pprn

ht a man ctivties chang the vironmnt the popultons in the lowrJes wi be moe datcally afcted tha ose na the smit . Tes atteeetons r eas tabe fo huma octon utzaton nd r mostdensey ocpied by B bpaala.

FUEE OF THE PHYSA EVROMET

To elucidte he nfluenc of environmta fato on the Cerro de Muee upn h distibuon d deni pa of B !bpdf4 aboroy we fed stdis were conduced on e moite d herm euiement

o the peces A eximnt anima were manained t rom emprtueof rom 18 o 22° e for not mo th wee po to expemeton Themetd for meung sol moire and mpee ave b pesentedearir In additon, reliab body ad mb sol·ar nterface empr weobtned fom soe 2 specmen

MSTURE REQU�S

Po to exmnon of eonmen moue reathip the tmoie conten of sx pecmns was detemnd. These niml wee paced

on sperstuatd filtr paper in cod galon rs for 18 hous nd ten dc·cted t 05 C fo 8 hour Tabe 9 gives petinent cod of hydrted


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/ E F T SUBPT IN T �'3

TAB 9

Tola/ wigb f B sulmata efor and a Juao.

ntl W. Dd c Wer Cno a G in G

8 2 76 2 26

8 0.33 2

2 2 6 62 2 80.8

CUNIGH (17) eoted amost he sme percetage o weight ·losfo sx esiccate &achoJep! paú/i1!. I animas wi iita weigts ro08 to 320 gms, he rortio o wate 15 wa om 9.7 to 85 % (806 %aveage) Te maest weighed oly 2 % o the argest yet the derence weght 1 etwee ese two a s than 2% Wate cotet was a essetia

y costat rooto o the tota weight id o basis o h ousion hatmaer smaers coti sigiiaty more boy wate ta large oesFe eerimet wee codcted to estabsh the citca eve o ater

os B !Nbpalmata. A 27 ams e hese eemets \ere eydtO sersaurate' te ae cose galo ja" o 8 hou A Theco rosn e (Moe 6) ws te to 40 C or 2 hors or to eeretatio to emoe ecs mse rom the abet ad te aowed t stabizeat rom temeae e isate abine! was rtte tertre to maitaie twee 22 24 C at a cotroed eate midy oom 0 to 8% Beoe eah weighg o a te baance au Cetog Mode ) he se was "wake oer absorbet aer to remoe xcess bod suEace wate. ae ws taken to aoid ay caca charge.eghts wee roe immeiatJ ror to xretato a at iteraso abot 4 to 60 mt ig he tests or at he ose! o ay obered strsSix cotro secimes were te ore moit chambe a sjetto othewse etca coditios.

o beaor attribtabe to strs was obsere in anmas hat a ¡stu to 23 % o ther iita body weght but deite aoae reato weobe wh te weigt s eceee ths e Noe o the smes sr

ied wate ss o greate ha 2% e atte ae s osstet wth that g by E 70) or Emati' eulz, bt ces te arage o47% i BarJep púicN CUNIGHAM, 7 o weght loss o sessress were oted mog the coo amas.

It was tcate hat bcase o saceoe atohs smaema wod se water ss moe raiy ta arge oes Ts heome n etei amhiians y E 70 an TORSB 7 Of


42/103

54 DE OPl

seprey esed B Jbpmat weighng from 130 o 4. gms the loss rae

pe u was fod ú b an almos cossen ¡nvse proporon o jni weghThe ress re prsened n Tale 10 Rs o oss e o 08 o 52 % erho avergng 5%

Cen behaio sposes unde s of wae wee od wonsancs ds aoomized he a he al we loss vel which sored ws no oserved Coling of ndvds s so common onc he seshhod is hed Animls sbe o dehyrion n he sme conne pssenly congreged n gh, b-ke msss whh efecvely redced heoa posed body surface area Ress from one xperme ine of heff of sh behvr is gven n Tbe A! s nmls "rved he 18

hour es o isod spemes ever srvvd eposre perodTe sma nmas pea o n somewha mQr advanag fom hs

copeve behvior hn age n Those wih nl weghs of less an gms hd he houry rae of loss edced y eas 09 as compared o gien in Tae and he averg houly a fo he peimelgops ws lowed from 5 o 11% a dference of 1.4% Simlr ehvorhs een oserved ding esvao of racoep púhu (CUNNNGHM7) and in prima ud on An,id" y ROENHL 63

TAB

10s ras B. sublm. dem/I·(ng agaio bcav EpeehJ bew d %

nita ecd Tim ost u/HrW W

30 88 625 32.3 26 24 6.2 28 4.0 2 62 262 2 .38 2 207 3996 164 800 16.; 202.0 88

80 69 08233 21 800 7.3 92. 23 800 6 4.71 4.36 80 74 0.9

ABE Wef u es o B suplta eO beho. et

ol e he s he aed Tbe

Intial Dicca % We os osW W of it W

187 6 199 69 82 1. 26 9 19.9 12.79 20 29 6 34 98 .


43/103

ECGY GSS SBPL¡ IN CS IC 5

TALF Reh)'drato 01 eyau B ubmt in tw mlos

2 / 4

N

il Dehyt iomt W vitoW Wt. H it Wt Re

1 1.99 9 Sbed 30 +0.; 1.2 46 1.97 b 2.8 04 7

2.79 20 er 26 -0. 95

4 .7 Mt 183 004 97.9 br5 0 7 Ms 96 97

Cbe6 �4 36 Mi 3 0. 96

b·

mmdiatly oowing deydraton to evels o betwen 15 and 33of nta body wet, a sers o six spemens weghng om 50 to 364 gmwr rehydrated Thee o ts wr sbmrgd n wate and t otr wer

paed n veed tn dih ind wih atuatd ltr paper Thes \e again maintained at tmars o bween 21 and ° C Prncnt murements on wate reory are rsented in Tab 2

hr meged mens avagd 9% o ntJ wgt atrte ors wile tose n te most ambr rganed an rag o 9 %Altog the dirne o te two aeag i onsderab (2% ) treeem o be e eooga adantage gand by ubmgene as omaed toaturatd raes n ates o eydration anma wr w aboe temoistre ts mit o 2% wtn te ors nd ad sentiay egaindte niia weght tog o bmrgd pmens weghed moe · hat ¡nita weight at this t it s sgnnt to note tat < 3 n eoering97% wa eded by wo o the most hambr pmns at 9 and 99%At he nd o e os a peimn qaled or ded t iital wegt

omparon o deydraton-ehydration ates o th me peimsannot b mad a agggton behavor dmontatd ding te om ttrted in modation o h weight los ra n only the most meanima are osidrd ower reoy ats mh greater tan deydraton at the m mpau aeagng 92 % o th oal t iod o 3 %r ho This i tme gte an te mot apid hoy rate 52%)

obsred in epaatey deydrated and twele tmes hat o tbeaeage oy ate o 5% COHE ) ung he sme aimas odeydraton and eydaion eermn ond AIees ¡ugb to se watr tme aste an t gined nd matila hscho/tú to lo timemore apidly t may be that h dernes ar d n pat to he that dehydaion ws arid ot at 2' C and hydration at 3 C

Athog t Ceo de a Mte i subjetd to ten a! dung


44/103

5 L C

mh of he year B ubp geney aoids anag ourss and nun·

atd aeas On only sx osons wre ndds seen n rely fooed miohabiat. Expemts we conduced o em he ength of tm es anmls an sve when sbmerg

3Sbmgec tolranc 01 B sup:mat p'us 2 J 3°C. �qH{s p

PL phyi% /

Sdad s, Wt. PL iference (gs) h PL (37.5 078 5 6. 259 ¿ 0 3.75 05 4 ¿ 09 7.75 15 4 50 . 4 . 4 ¿ 054 64 4.0 .6 < 55 31 < 11005

1.99 < 00

Pin ja oaining 300 m of wel wa wee allowe sabze o24 hous a 22 o 23' e beor epermnaon. The wae was no erae ig e eperiment Animals we manand a oom mptus n cntan- pay lled wh moss fom Ihe hta se and tesed wihn a wk o pre Tab 3 inluds th obseraons on e epmna anmas. e onolgoup aga ompse s spemens mantaned n coved most ambesg expimenan. Lo% f ighng abliy, (capr thrhol) eiherola or in respons o stong essue ppled o he hnd lmbs

epesents he lm byond whh escape s impossl Te physologiallehal pont w aen as Ie essaon o apllay flow n Ihe gargio. s cra ae snally Ios use by S (70) Timnevls are gin o he neast qarter h! drminaton Losf ging abliy btween .50 and 7 hous wt anavage o 5 A muh br ange o olen was demonsaed physologcal lal pons 675 .0 hous; avage 10 ) . Des eos bewen CT an aeag 30 hous range 2 o 60 hour) No bhaioral or phyiologca repnses wee obr n he conrol go. Myinoman is naea o usty mo hn a geeal inpan o holranes t s edent howeer tha at gh hous bp h suferevsbe lethal amage NS 70) oun n nvs oeon bweense an oranc o sbmegene in E ho whh no ppaenn Is experm xim me for oss of ighng abty ( a"logial m of NS in eogaphc ces of Ew was 9


45/103


46/103

58 BIOLO TOPC

spemens in he 37' e room wr dead Wn aced n a pai pan nanng

5 cm deh of water rehated to 37° e five oh B Sp"t isanlrespnded by vt thrashng f bod nd ai dng frm therma ;hockwthn 3 sconds.

ROSENTHA 63) observed that d Igr dd not srvie moIhan 24 to 4 hors afer exosur to _6 e fr 5 6 mines and ondedIha tmperatrs mh elow frig for more than 3 minutes re fatal Term-al tress n And was demad at an average f 3 e f! eght nmasad the riia therma mxmum (C) indcaed by t onst of spmswas aained a aboe 33 C In xpriments wh En z; TBBN7) obaine sress resnses

a

t6

C and a CTM f3

t3

C.ZWF

8) and UTO 4) have reored crreaons f C ad habadiffernes in sveral speies f salamandrs Of plethoontid species andsbspecies nded n UTCJO' sudy, he averags of T's for adultsamated a 2 o 2 e wer from 335 e for Gyrnop 4 d) to378 C (for Mu qrdii. Conger speies suay demonsratd smar eTs with th expn f gn' whch ebed sg-nfn diHerenes btwen D qw 3359 e) nd D j j362 e).

Cloaca mperaures fr spemens of B obaed i the

fid osely approximated the iar interae temperatr f the mirohabiatFg 35) . Body mperaures w en mmdaey afer ncoerng h an-m in its rfug. The hnd foo w hed d th thermmeer nsered and eadat s stbilizaion point wihn 15 to 3 sods f he pemen xhibited markedactvj or the hermmer d sabilize befre 3 secndsJ he reading asdisregrdd. The sma siz f vnles dd no prm body temprare to btaken whot mearable h ndct from the obsev After rmvgmoisre frm the thermomtr ip the soiír erfae ws then sred bypressng the bulb io the ssaum so at ony th · upprmot srfae was

expod lhgh varius factos sch as behair prios exposre, variatiosin dn from th subsraa and evapratn of bd mosure an measur-ably infne body empratr f amphibas, i pleodontid derss s usuaUy oe he tmerare o e mcrohabia (Rfr t AT_OM, 1 for a general re) ha be oudd h the approximaiof bdy ad mbent tmrars is a resut he nessi fr these eresratoherm t ann bod msure b clse ona wh th wt sbsum(G ; ATO ; eON 6) .

Ove 6 mpeatur tke of microhabitts ied by B f

have ranged from

3.8 C Sh eremes ar aaned ony nfreqnlo e Crr and hgh tolean of lower hrml evels, at whichit is no normaly ate the specis is rare fund at ambent emperares ess of C. The range f body tmperares at whch aive anm s habeen obsed is esmad frm radngs ten fr refuga a me of emergece 188 ative anma my erds rage fm 6.4 128 e aerag88 ± e) \ ) offers n exensve ummay of "vlntary


47/103


48/103

VSA I

TABLE 14

Geo¡rp, DstribuÍ01J o Phooi G i /h New Wo

. , < • B • "Ñ i > J ( • v

og( XLuogh Xh�ogh; X

Aides X XBrcoJ�pJ X XEtn X XHmdylu XId XSeoJ X

EUf)'c XG)"zoiu XdOr XHd( X

nuu XPsoio XTyp% o XTh'i X

/goI X X X X X X X X X X X X X XChipeo X X X XOedPia X X X X X X Xr-/limg X XPdea X Xo1 X

Thous XHg


49/103

ECOLY OP BOLITOGlSSA SUlM IN COA RC

TLE Il

AJ habÍl 01 Nw Wor xt/opi p/hodonid ga

Subfily Sam T, RockGUJ Pnd tr Bn \X'o be

DsgnhDsmogh X X XfOgh X XPhogh X

Pethodntn

GyiophiI X Xcudorio X SlohiJ X XE X X Xnu X XTphorio X

Hd6rio XTphomo XHidi. X Xhdn X X XAd X XEna XHdo(¡ X XBho X

TBLE 6

Hi di,; o roi 0 phodod d8r

Gn

BoiooChroprodipin!iogdourLnioTo

Sfee

XX

XX

wi

Sxs

XX

61

A

X

rbe

X


50/103

62 RIA D OOGA O

species known Erom lowand o subalpine regions Nine genera inude peies

wh prmariy aquati or sreamdwelng adults (Tabe 15 The most extemeamps of aquati lie are two lte known monotpi geera found in caesand artn wels he apparety paedomorph Hidoi and TypoogeSevera spe of Ee al aan s mary wthout memorphosyploion s not kown to depa rom its underground aquatic hbtat; whieLeuognu oony upes sea beds sedom emerging byond eposerocks. Seeocu res sowmovng strea O ond in ow eeationsGynoilus and Pdorn mark a trastion bewn aqtic ad terrstrialhabtats Aduts of thee wo gera are sem an'd seam bak dwells

The widt habta uiizaton is exhibited witin he gera Do-u and u Both inlude species of aquatic strem bank or woodndhabs. l s somewhat moe teral hat mot seces of '&Hedyl is known rom both srea banks and woodands, whe Peodo Wqutoably a terrtral genus only ne spees P dU, s am bak orm osp Ei and Peog are wodland habitants wth soe propensity or brwng A specal type o terrestrial habtatthe rocky slope s tlzed by oe spes o Aeid, Hdo and Pleo·d; gera most ypialy rom woodland. It appears that e ¡aoous hbiths been dered from a more generl tereia adpation rather an through

dr tson rom a aquatc stence, as the oy genera t inlude rckdweling spei are those wth terretia or wodand reprsentatives.

Modern tropil plethodontids occupy considerabe range of habtap but no known tropica spei an be ecoogicay equtd with the aquaticor s bak dweles o tempate attuds Jt seems probable that since dirteestri developent has emerged a a broad adaptaton there h been lselee adage n returning to water Diversiy has reslted m efinementso eensons) o tilizaton o the tertra envronment for whch dr embyQ-10ga development ha bn a prdptaton.

Most tropi samanders are o a genera trrestra habt ad re groundwndere living under cover provded by ler onheles within he severeogized genera omtting the igmai genus Hpog) species manstvaring degrees o speciazation A smary oE the more ditinve habits igen n Tabe 1 he restrcted abs kow or soe genera e.g. PvioJge,us ad rio are undoubtdy a rst o lited coeing and n·adequate knowedge of tropical saamander ology Oedipi1 and Liioare elongate� worm-lke fos wel adaptd to a ubterranean exstence ausops nd rok ubble are especiay Erequented by opo


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ECOLY OP BOTOGOSS UATA CO R 63

brmlads o aroids hang scoop_shapd las wt bass tha sr as srois of n watr. 'eca xen of "aa wamp sy depndpn te compeiy o h or and may tend ben about 0 o 30 m insord jungs Most rport of bora abits in ropal salders inuderefrnce o bromeliad dwr. Crtan spi, Chrptri pd Cif al and Blit(l ald, ar known ony from this habta

Te gratst gnrc diversy n habtat utlzation s own by h-rrM and BIga Wthn later genu no specs ceeds t valnceof B(l( pa/ whih reains h unspced errsria rof he my and ehbts a varey o abts (Table 6).

Te extent to whc e hstora nd prnt dsrbuo o samadrs en influced by mpraur ad avaal mostr cannot be ovrsmaed. cause o he inimae oogica rlaionships of he wo aos is desirabe to consder m togtr t is orunat tat an insrent su as tOD (39) mod s avlab by wh ts rlatonshp can b mpiraly defind

Consn wit my prvos use of ts ytm n reng o te oldstbuton o Blgla bpa rcnt cgograpc pats plodonids and possiby aU saamandr an b masd in m o bimperatredicussed ier and h ponia evaporas?iraon ra oa eapo·ranspraion (PET) is he amount o watr lo b vaporation and rnspirationprsumíng watr be ontuousy aailable T B) K wr B s !man annua bomrature and K is a consan vau f 58 To oba !hennua rae o PET pontal vaporanspraton dded by te m anuaprcpation (OD 4, 42)

Supeimpoed on h be dig of Hodridge ssaon oOim Vgaion ormaions peed n Fi 6, is he gea ecoggrapc disrbution o pltdontd amand�¡ h attdna regons oc·pid by t amy ar Tropcal, Low Subtropil Warm Tmpae ad Cool

mpra. quaed wih hse rgions ae Topcal Subropca Low Mone and Monn a1dna bls Eac lattdina gon deed by bompeau ts nludes egaton omatons ndiad by ah o t xagonswhi !a aiude rgio d a of !ose owe biompeaes. Fo ampa oa of 8 Tropic aidina formios ae defined conng s afo Wam mpat and ow Subtopca wh are spaatd by a rtcatmpratu lne o 17 C) T sn atdna rgon inud a total of 30fQmaions. ou egios and 30 formaons ae by lehodoni whare strcd o !h Hmd Phmid and Superhmd Hmdiy Provncidcatd a h bas of Fg 36. It woud b a mtake to asume at a neatad ar-c disibuona piue saped rom he pcie mod o Holdidgxagons ponids Salamadr and many o!her animas ehiehavoral and physiologc adaptatos provdng a means o cape romdrec pou[e to pysa nvromnt. Fr ampl, brnaton aestvatond migion are ma o vading faorab erm and permi biaon o aa othwse cologcaly inaccsibe


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64 RVSTA DE BIOLOGA TOPC

I brad perspetive he dstrbuto o pethodontds is assed th

o recogzbe ms. Serios moise resricions ae met whe he PEro approimates 100; n other words, whe PE eeeds he men nulprepao. Uuestonably the .00 mosre ine s no mitng fto fospees apble of voidg droht odtons. The d fores o he SubhdPovine re margna hbis wh may be vded by wy of more heviets. A exen exmpe o he opton of mginl odo isdemonstrated by soe samnders of e warm empere regon in he weenited State, sh s Barhoseps a/a, Ans gb nd Ehht tht nhb he ek ores ommuy o Caior Although auly prt o e Wrm Temperte Dy Fores ormao he Sbhud H

mdy Provnce his couy is cped by hese hree spees in locedeas where opogrphy nd ange ompesate for cimaicy indeqemostre

ew ohr amnde hbit he tides t which B S1bpm isooly foud Poy ,ex s budn in he pne forest belt of Gt-emala the Topcl Monane ltitdl bel (the co empete ae oS 65) Norh Amer AI hd and H)'ozs pla-"phaS are hgh alttude es ocurrng dfee loclies t mOre h2,700 m bu eher o hese spee occpes the subpi bel Te pper he-a bundry o he Cool Tempere lattudinl rego o he Motne lttudnbelt o 6' C bioemperre is probaby ne the mmum lmt fo slamandershe summt are of he Cerro de Mure s maclly Tropical Monaebe

By sng the 6 boemperre s e mmm herm lm pleho-donid dsbion nd ppying he metri mits of ttdina be give by. HOLDRDGE (39 2), the mamm eevao ailable or hbs ould be

bot 4,000 m on the het eqo titdina inses w edúce this limoncomtat wi owerg of he Moae_Subpne boudary. The re ooaly records by whh o verify the once of lmders t �his esmed

mxmum eleato n opc ones, thogh Chirripó Grade (,820 m), e souer Talmanas of Costa Rca, ofers a opporuy o s hs ide.lamader re ot cpable o prolonged epsure to sbhumid codos.

As a rue moisure ad tmpeature operae together in lmtg he drbon terrra speies. here re, however ertaj codtons in whh he oppear to opeae ndepedeny la thema «gons caaestll oped bya spees moste exets a determinng nec Fo eampe Ad Jg.bi o the warm tempere egion Calonia ehbts a clse assato wihsoi mose nd mon of prepttio whh ROSENTHAL 63) consders o be more mportnt hn tempertre.

The humd provc are sbeted o herm ves which y imtsalamnder dstrbuton. B subpJa s esealy lmtd o Lowe Mted Mone therml levels wth he etensve Sbrop bt of mpbe hmdy The presen liudnl dstrbio of tropa pehdotdsis evdene of dption to erma evels wihi the huid eions Mos rp


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EO"OF COSTA R : 5

pes r und forms reeig hei og om e l tempeae OldNohe hepetof (SVG 64) Cerai geera, mo oaby BoiogloJ!d Odn" hv been ésfu in opyg he mois owads.

e e distit diffeee i he yps of effes of eionmeatmerars pon B bplma Mh of te work do o the r thrmamma (C of exrropia amande h be revwd by HUTCN'ON Howeve evdene obned duing h peent dy indes hae lueof C may be of ght ologcal mpotace For eme he Co B m li somewhre ea 3' C y he pec nnot toee

polged epsue to 27 E me importan is the herm eel ingspees distribuio whh mor aey refes he nre of adapto Thewide nge of ouy tmpeaes eoded n he micohabit 1 C)in9ca broad eologal orance n adl, b ther ar be, ye cria!aspes o the pre. Mohbas my hve emerres abve 20 C a timesbu B /bm dos no m to be pab of ned bdng behviorhen bjtd to proongd pose o s herma ves Moreovr the eeogil ermal mximm s probably e bove whih egs hve be bseved suive "tiued xpsue

Eoogl herm li pt are rpngly mch belowtho of Dog1t jUSC, whose eg ZWEIFE (8 hs ssfly -bated 26 C It shod be noed howvr, ha he iudin dirbio B lm is generay evale n bomperare o he cool mpratdi\a regons opard to he m mperae egio oped by D. 'j heefoe he opi om h oe thma limi hn hoe oed byhs ext�pca peies.

Pedod poduive ptes lified ino hee bas pswih oy t overap. I bu óe ges' th fe s e noably onsitend be tegoized he foowig manne. n Type the most pmitive,he eggs re nhord i ar nd an qua arva psesng fnonl gilhatces pon rasfoaon, he dt om is tpaUy i emiqiTye II iudes forms whe ggs are deposied o d a hve rvae amigte o wa. Afte meamophois he adul emeg o upy a teresihabiat n Typ III compete errsram s hbd nembonaed ee depositd lard and undego die deopmet. The mbyo hathes s rpia of the du and oes h lao gls daely pon 'hathng osoon frwrds A sry of h orne of produiv ye n mreo gener s gv n Tbe 1

Pbably he odes and mos genelzed vig pletodoid gens isDeImognathJ (DUN, 22; .WAKE1 77) wh ao nne ecognzd pecie . Inhis grup al of te paers jus decibd e reprsentd. D.

t

r

dc¡a

IH

s n qi form ' a11 phas of s e ye. Trrtrial eggs nd dlt sg�e psen j D. fus/, bu te lave et water upo hting d do o emrge unt mmorphos. D. wrighli is trresal in aU pha; the egs undc-goi die deeopmen o ad


54/103

ISTA DE BOL !PJU_

u of Ihe occue 1 rpod"v pes ;n e zne g 0 odo.d, Th b 01 nued phs) eah ge si ha 0

\'ake S ex fo depon 1 haCJ

1

DeJ:oghJ LeuroglatbJ )eohilJ ( )U ( 0)Gyrophi ()phlwon 1)Ps

03 vial bolitoglossa

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