MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 160 The Life History, Social Organization, and Parental Behavior of Hyla rosenbergi Boulenger, a Nest-Building Gladiator Frog Arnold G. Kluge Museum of Zoology and Department of Ecology and Evolutionary Biology The University of Michigan Ann Arbor, Michigan 48109 Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN June 10, 1981 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan 48109. MISCELLANEOIJS PlJBLICATIONS MUSEUM OF ZOOLOGY, IJNIVERSITY OF MICHIGAN NO. 160 The Life History, Social Organization, and Parental Behavior of Hyla rosenbergi Boulenger, a Nest-Building Gladiator Frog Arnold G. Kluge Museum of Zoology and Department of Ecology and Evolutionary Biology The IJniversity of Michigan Anrl Arbor, Michigan 48109 Ann Arbor MIJSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN June 10, 1981 CONTENTS PAGE INTRODUCT'ION ............................................................. 1 MATERIALS AND METHODS ................................................2 O~igin01 Samples ......................................................... 2 ODSS Variables ........................................................... 5 St:~tistics .................................................................. 7 It~cliviclual Marking Tec hniqucs .............................................7 Size Mcasurcmc~lts......................................................... 8 THE ORISPO DIVERSION STIJDY SITE ...................................... 9 RAINFALI . AND FIAOODING................................................ 11 Gcnel-al Charat tel-istics .................................................... 11 Correla~rsof UI-ccding Activity ............................................. 15 ADIJ1.T CIIAKAC'I'EK1S~I'IC:S ................................................ I!) Color and Pattern ........................................................19 Prepollical Spine ......................................................... 21 Size ...................................................................... 22 GI-owth .................................................................. 27 Food IIabits and Far Bodic ............................................... 29 Clutch Frequency .........................................................30 Cluttll Si~e............................................................... 32 Egg Si~c................................................................. 35 Time of Ovulation ....................................................... 36 TIIE NESI' ..................................................................37 Materials and C:onstructiot~ ................................................37 Nest Dynamics ........................................................... 42 SIJRADI1I.T KIOL.O(.Y ...................................................... 53 . Ovipoution .............................................................. 53 1'1~crnetarnorphic-Crowtll i\nd Maturation .................................. 55 Metarrlol-phosis ........................................................... 61 Postmetamorphic Growth ant1 Marut ation ..................................ti5 Subaclult Survivorship ..................................................... 66 The Ad;iptive Signiticarlcc. Ot The Nest ....................................73 ADIJL'I' POI'IILATION BIOI.OGY ........................................... 75 Density Anti Movc*rncnt Among AI-cas ...................................... 75 Density And Movement Within An Area ...................................78 Length Of Kesitlency ..................................................... 86 Predators ................................................................. 88 Daily Activity I'attern ..................................................... 90 SOC:If\L KEHAVIOR ......................................................... 93 Bchavlot-a1 Diffcrcnccr Among M;rlcs ....................................... 93 Vocali~ation.............................................................. 96 Atlvertise~ncnr call .................................................... 96 (:ottr tship (all ........................................................ !I9 l'erritorial call ...................................................... 100 Encounter calls ...................................................... 100 Agg1.c. saion .............................................................. I02 Pail- I;o~-mation.......................................................... I11 Paternal Care ............................................................ 116 MATING SYSTEM .......................................................... 125 Matir~gSuccrs\ .......................................................... 125 Intensity Of Sc,xr~alSe1c.c tlon ............................................. 130 Nightly Sex Ratios ...................................................... 1.35 FI<;IJIIE I 1 Di;rgrarnatic actial view 01 the ODSS wlletc rrlosr of the ro.crtltj(~r,gr ~esearchwas conductctl ........................................... 2 Diagtarnatic aerial vicw of the water course along the rilain ro,5~)1brrgi study area labeled A in Fig. 1.. ................................... 3 View ot the tnait~pool looking south lronr stakr 0 in 1978 ......... 4 Vicw ol the ~)t.inl;~l.ysectiorl of the tn;iin ro.trr~hrrg~study area looking south from stake 150' in 1977 ..................................... 5 Avct-age rnonrl~ly rainl;rll recottird :I[ the I>as C:asc;rd:~s rnetrorologic;~I station plottetl according to ycar (prc-1977, 1!)77 anti 1978). ......... 6 C:~~rnulariveaverage montllly I-;rinfallrccortls obtainctl at tile I.as Castada\ rr~rteorologic:rl statioti plotted accol-cling to year (prc-1977, 1977 ;rnd 1978) ............................................................ 7 'l'otal numbers ol (lays during tllc wrt season (ilptil-Novc.nll)cr) with rairllall 2 2.5 mrn plottctl accordirrg to holrr of clay ................ Incrcascs in warcr level ol tht. main pool ~)lottctlaccording to the coincident rai111;rll tnc;lsl~tedat thr I.as <;asc;rtlas rnc,tcor-ological station 'I.l..icing : of the actual I-irt. and f;111 of the watct level I-ccortlrd on July 20-21, 1978 ...................................................... Numbers of ro.\(,nh(,rgl tn;ilrs obsrrved each night (July 5-August 22, 1977) plottcd accortlir~g to the amount 01 ~.;rin tll;~t lcll duting thr prccccling 0600-1 800 211 ........................................... I)atcs ol TOJP~~!)CTSL ;~rnpl(~xes...................................... I~orsolatctal v1c.w of at1 ;~tiult rnalc r~~rnljrrglcollt.c.tc.tl at Surnn~it C;ard?ns, <:anal Zone in 1976.. .................................... Positive print ol a 1atliogt;lph of tl~.l1c;itI and lorelirrrh~ol ;ltr adult nralc rosrnbrrgl ....................................................... F.nlat-gcrrlent ol tllc- left 11;111dof th(. rr1;lle ro.\rnbrrg~illustt;rtc~ti in Fig. 13 Vicw of ;I left p~rpollic;rispirlc, rcrnovrcl [I-orn;i cl~rrrlt.sritlI)cctlrl)rc~l,;~~~(~(i skclcton of ;in adult rnalr ros(>nDpr,ylcollected on I'ita Ko:rtl, C;;rn;~l Zone ill 1!)7(i .......................................................... Vcnttal view 01 thc riglit h;~nclof 2111 adult tnalc TO.\(,TI/~PT$~\11owitig 111c fleshy sllc.;~tlrrtrc;~sillg the ~)rtyollicalpine (fro111 Kluge, 1979) ...... IIistograrn ol live rnalt~ant1 f(.nlalr ro.\cnhrrgl W'I' and SVL. ....... Live rosr,rrhc>rg7 rnalc W.1' plotted arc-ording to S\'I. ................ 1,ivc gravid ro.trr111rrg1 Mr1' plotted accot-(ling to SVI. ............... Livcx Icrr~alcro.crnhergl WrI' attcr ovipos~tionplotted ;it cortling to M'-1' I)eforr \lt;~wnirlg ................................................. Clla~igrit1 ro.srnDrrg-I W'I' and SVI. per day.. ..................... Days 1)etwecm \~~cccssivcclutc Ires 1;rid by fcrn;:lc ro.srnhrr~q~.......... 23 C:llrreh rim (rlurnbel- of eggs laid) of (lost,ly tclated sl)ccic ol xladiator flogs, hoan.c, (rrp~t(~n.\irr~d ro.\rrzhrrg~, pIottr(1 accot-dittg to SVL .... 21 Seasonal tlutc.lr six vzuia~ionin indi\ritlnal lcrrl;rlc rosellhrrgz ....... 25 Diagtarn;ltic prolile of a typical ro.\rn/jrrgi nest at~tlits tcsid(~t~tc:rlling 26 Male ro.\r~~O~rg~110. 144 xivit~gthe atlvertiscmcnt call from nest no. 127 on Aug~tst1.1, 1978 .................................................. 27 A ro.crnh(v,yl r~c-stwit11 high rnud ramparts at Bal/;rpatril)a, Ecc~atlor . 28 I,ocation\ of all ~osrn/jc~r~qir~csls