aa Spek EE ee ‘Comparative Grouping Behavior of Juvenile Ctenosaurs and Iguanas Jost M. MozA, Department of Wildlife and Fisheries Sc- ences, Teas AEM University, College Station, Texas 77843, usa! MeNab (1963) suggested that in species with low ‘competition for food, grouping can be advantageous. Herbivorous species generally have smaller home ranges than carnivorous species. probably because of ‘the greater concentration of food resources within @ given area. I this reflects reduced competition for food resources, then grouping may be more itkely to ‘ocear in herbivorous species (McNab, 1963). Burg- harde (1977) and Burghardt etal. (1977) documented ‘a high degree of grouping among hatchling iguanas, ‘Iguana iguana, herbivorous species (Randet al, 1990), during departure from the nest. However, little i= mown about the degree of grouping in juvenile ‘ctenosaurs, Ctenosaurt similis, another communally ‘nesting species (Mor, 19892), Because juvenile ceno- aur sre mainly incectivorous (More, 1986), com- ‘Petition for food may have been important in their ‘evolutionary history. The degree to which hatchlings ‘and juveniles are found in aesociation may reflect the level of intraspecific competition. To evaluate this ‘hypothesis, [observed hatchling ctenosaursand igua- nas leaving nests in the fold, and behavior of jave- niles in enclosures. Tobscrved ctenosaurs and iguanas in the Rafaol [Lucas Rodriguez Caballero National Wildlife Refuge (Palo Verde), in 1985 and 1986. This area ‘extends over 7354 ha in the middle reaches of the ‘Tempisque River, Guanacaste Province northwestem, (Costa Rica, (I02I'N; 85°21'W; Boza and Mendoza, 41981). Iguanas and ctenosaurs share nest sites in Palo Verde (Mora, 1989). Tobserved 181 ctenosaurs from 51 nests, and 69 Aguanas from 15 nests in the field. More ctenosaur nests were found because ctenosaur density is higher than iguana density at Palo Verde (Mora, 1988). Al- ‘hough ctenosaurs and iguanas can emerge at night (Mora, 1989) all eld observations were conducted ‘between 0700 and 0900 h. When T found a nest with ‘emerging animals I remained at a distance of 20-30 ‘mand waited until the next emerging animal ap- ‘peared at the exit to begin the observation period. ‘The time an animal remained atan ext beforeleaving. land the intervals when no individuals were at the ‘ext were recorded. Taso recorded behavior at time fof exit. chose these rs because mean time at ext could reflect the degree of vigilance for pred- ‘tors, and mean time between animals could reflect the degree of association among hatchlings leaving the nest Behavior may reflect the degree of vulner~ ability to predation. Time was meacured tothe nearest (05 min. A nest was abandoned and another sought if 20min passed without another animal emerging- For the enclosure experiments the same treatment setup was used for 40 iguanas and 40 ctenosaurs in ‘each of 2 years. Hatchlings were housed in three en- ‘looures of 15 % 3.5 «35 m. Bach enclosure had NOTES natal ¥ ‘and one wood platform (1 x 1 > i'm) where animale could bask and the ‘The enclosures also included six bamboo tubes that animals could use for retreat, rest and thermoregu- lation. Both availability of sites forzetreat and density of individuals can influence the number of animals ting alone. T that the number of animals eating slone wee dereaein enclosaes with ew cr sites for retreat or higher density of individuals. ‘To assess this hypothesis { housed the animale under duce different combinations of density and number of retreat tubes: 10 ‘and six bamboo tubes (enclosure A); 10 hatchlings and 12 tubes (enclosare '); and 20 hatchlings and 12 tubes (enclosure C). ‘As resting group size may change with lizard age (Barghardtand Rand, 1985), collected data only from {young animals (0 to 4 months old), and used a dif- ferent group in the second year. I visited the enclo- sures at night (2100 h) twice a month for four months fn both years, and recorded the frequencies of dif- ferent group sizesin each enclosure. Hatchlings more than 25 em apart body to body) were not considered to be resting together. ‘analyzed all data using nonperametric procedures (Conover, 1980) Chi-square tests were performed on ‘the field data to test forspecies differences in (a) group size leaving the nest, (6) behavior leaving the nest, fand (@) numbers of ctenosaurs and iguanas resting ‘lonein the enclosures. For each species, a Friedman test (F) was conducted to test for treatment effects on ‘the number of animals seen resting lone in the three enclosures used. “The N given for the Friedman tests is the total ‘number of animals observed in these enclosures. ‘However, the degrees of freedom are given by the number of blocks (nights of observation), and the ‘umber of treatments (enclosures). The percentage of ‘solitary animals, the time animals remained atan exit, ‘and the interval time between animals were com- ‘pared between species by means of a Mann-Whitney Deeet TA). After emergence, 122 of 131 ctenosaurs ran from the nest toward vegetation; in contrast, ll iguanas cbacrved walked slowly in groupe from their nest O@ Mas, P < 0.001, N= 190). ‘The mean time animals remained atan exit was 280 1500-3541, N= 131) for clenosaurs, and 52 (0-270, N = 69) for iguanas; the mean time without animals ‘at the ext was 80 (0-420, N ~ 80) and 03 s (0-20, N 50), respectively. The mean time animals remained ‘tan ext and the mean time between animals exiting, Giffered significantly between species (U= 5.94, P< 0.001, N= 200; and U~ 3.68, P < 0.001, N= 130, respectively). In the enclosure experiments almost all ctenosaurs were resting alone and most of the jguanss were in ‘groups (x2 = 235, P < 0.001, N = 628; Fig. 1B). The enclosure treatments did not significantly affect the age of clenosaurs resting alone (F= 1;df—~ 2, MézP > 025; N = 340) or iguanas resting alone (F = 1.643; af = 2, 14; P > 025; N= 280). Overall, the percentage of ctenosaurs resting alone was signifi ‘antly larger than the percentage of iguanas resting alone (U = ~3.508, P < 0.001, N = 24).NOTES 338 +90 B x0" Bl croswuRs = Bi euaws 20 18 10 1203 4 8 6 7 oT GROUP SIZE RESTING Rig. 1. Frequency of group sizes for ctenosaurs and iguanas leaving the nests in the field (A), and esting in the enclosures (8) In this study hatchling and juvenile iguanas were highly social animals, thus corroborating. previous reports (Burghardt, 1977; Burghardt etal., 1977; Wer- her etal, 1987; Burghardt and Rand, 1985). The anal- ysea presented here indicate that iguanas did not walt gat the ext, and tended toleave in groups, where as clenosaurs paused individually atthe exit longer and then laft one ata time (Fig. 1A). Although Mora (1989) found 18 hatchling ctenossure at 2 cm from ‘proximity in the wild, and I never observed fwo such Individuals ‘lenosaurs sleeping close together in exptivity (Fig 1B), but I never observed a pai resting Jur Ing the day. The different enclosure conditions did aot affect the numbers of animals resting alone. The difference between the two species does not seem t0 bea response to either availability of sites for retreat or density of individuals. However, more treatments with higher densities and/or manipulation of food 285 might be setup to compare the degree of intraspecific ‘competition. Ctenosaure spent more time than igua- tas af the mest exit before emerging fully. This is Consistent with the hypothesis that they are looking forandavoiding predators because they donot emerge from nests simultaneously as do iguanas (Burghardt, 1077), Other factors could contribute to differences in susceptibility to predators; eg, hatchling ctencsaurs fare smaller than hatchling iguanas. Ctenosaurs are pale grayish brown with darker brown marks at [batching (Mora, 1980), wheress iguanas are green. On Dare ground, like aexs surrounding nest sites (Mora and Chaves, 1969), clenosaurs are probably more ‘cryptic than are iguanas. For the predatory ctenosaurs, relatively high intra- specific competition may select for reduced associa- ton among individuals. Thus for hatchling and ju- ‘venile ctenosaurs the best anti-predator strategy may be to simply run from the nest hole and seek cover. Further observations on ctenoseur and iguana behav= sor when emerging and leaving nests, and subsequent movements, ane needed to determine the effect of ‘on the behaviors of these two species. The ‘ofa blind in sites of shared nests would {aalitate such observations for that objective. More {information on any Hind of ctenosturiguana inter ‘action in those sites is meaded, as well as more about ‘behavior of hatehling ctenosaus, che les-studied of the two species. Adda —This research was supported by the Eocucla de Ciencias Ambientales, Universidad Nacional, Costa Rica; the World Wildlife Funds the Organization fer Tropical Studies: and the Subdirec- cdgn de Vida Silvestre of Costa Rica. J. M. Packard ‘made important ‘for the preparation of ‘the manuecript. I thank C. Burandt, C. MeDonough, ‘and. Rasmussen for their critical review of the mant- Script Special acknowledgment to D. L Werner for her idess, hospitality and friendship during my visits to her “Iguana Management Project” in Panama. Lrearume CreD BozA, M, AND R. Minpoza. 1981. Costa Rican Na- ‘Gonal Pazks, INCAFO, Madrid. 310 pp. Burctasot, G.M, 1977, OF iguanas and dinosaurs: ‘socal behavior and communication in neonate reptiles. Amer. Zool. 17(1):177-190. SETH. W. Grain, AND A.S. RAND. 1977. Social ‘behavior in hatchling green iguanas: life ata rep- tile rookery. Science 195(4279): 689-691. Jano A'S. Rano. 1985. Groupsizeand growth rate in hatchling green iguanas (iguana iguanc). Behav. Erol. Socobiol. 18(1)-101-108, CConoven, W. J. 1980. Practical Nonparametric Sta- ‘istics, 2nd ed. John Wiley and Sons, New York. Sopp. McNab. K, 1968, Bioenerpetin and the deter tminidon cfhome ranges Amer Nats S730 ery Mota] Me 1986, Alimentactin y cecinient co- orl del garobe, Clecaur ss Gry, en 50 Primer ate de vida. Uapubl MS. These. Univ. Coa ie San Tose. 9p. A aobe. Ece-bchvioal espe of two com- Tmonally nesting iguanines and the stuctae of286 their shared nesting burrows. Herpetologica 45(3): 293-298, 719698. Ceensaure sims (Spinytaled igua- ra). Nest emergence. Herp. Review 20(2)53. 1550. ‘an instance of albinism in the cteno- ‘aur lizard, Ctenosaura sis, in Casta Ric. Bull. Chicago Herp. Soc. 25(6)70. Sat AC. Caves. 1989. Laxacems bicolor (Gurrowing python). Size. Herp. Review 20(3)72. Rano, A.S. B.A. DUGAN, H. MONTEZ, ANDD. VIANDA. "1990. The diet of a generalized folivore: Iguana iguana in Panama. J. Herpetol. 2(2)211-214 ‘War, D. 1, EM. BAXGR, & Dat C. GONZALEZ, AND TLR Sosa. 1987. Kinship recognition and group- Ing in hatchling green iguanas. Behav. Ecol. So- sobio. 21(2}:85-88. Accepted: 16 November 1990. Irena of agloy Vol 75 No-2 p26 28, 191 1901 Soy forthe Sta 8 Amphibians aa Rpts Juvenile Flatback Turtles in Proximity to Coastal Nesting Islands in the Great Barrier Reef Province TA Wau, of Enoronment and Heritage, P.O. Bex 5391, Toonsoile, Queensland 4810, Australia. Sea turtle hatchlings are widely considered to dis- ‘perse to the ocean where they live apelagicexistence for their early years (Brongersma, 1972; Carr, 1986). ‘A pelagic habitat seems likely for juveniles ofall spe ‘ies except those of the Australian latback turtle, No- {olor depresa. The Presence of juvenile N. depresst ‘yemnants at feeding stations ofthe white-bellied ea tagle (Haliaentus Iecegastr) on Great Barrier Reef is- lands was interpreted as evidence for the absence of _apelagie phase inthe lifecycle of thisspecies (Walker and Parmenter, 1990). The captured turles ranged in ‘carapace length from 14 to 24 cm and were located ‘oninshoreislandsalmost entirely at the southern end. (Gi>-24) ofthe Great Barrier Reef province (S22). Following the previous report another 45 juvenile turlle remnants were located on Great Barrier Reef {lands in December, 1959, All except one of these ‘were recovered from Arch Rock (25°22, 150°57'F) ‘and Cullen Island (21°25, 49°29). Arch Rock is small, steep, barely vegetated zock 17 km S of the prinepal east coast atback turtle colony at Peak Is- and (Limpus etal, 1983).Skeletal remains of 19 small turtles were recovered from the top of Arch Rock. Material consisted primarily of hyoplastron, hypo- plastron, costal bones and an uneaten head. Callen Ibland isa small grasy and shrubby island 64-74 km [Nof the major east coast colonies at Aveid Island and Wild Duck Island (Limpus etal, 1983). Four juvenile 1N, depressa shells and skeletal remnants of 21 bodies ‘were located at eagle feeding stations dispersed around Callen Island. (Curved carapace lengths of decomposed juvenile turtles were estimated from comparison of hyoplas- ‘on and hypoplastron bones with those in intact ear NOTES a8 Numbers of wont tuts ‘al libel we a 4 i 6 7 8 19 20 ot ‘Carapace length (em) Fic. 1. Size distributions of turtle proy remnants collected from Cullen Island (black) and Arch Rock (shaded) in December 1989. Curved carapace lengths fare in most cases derived from matching of hyopias- ‘zon and hypoplastron bone sizes with those of intact carapaces. _apaces of other specimens. The size frequency distr- Dation of turtles preyed upon at the two islands is shown in Fig. 1. Carapace lengths were 129-205 mm (= 175 + 19 mm) at Arch Rock, and 113-197 mm 164 22mm) at Collen leland. Thespecimens from ‘Arch Rock were slightly larger than these at Cullen Island but the difference was not significant (t= 1.88, P~007,N= 4d), The similar body sizes suggest that ‘eagles preyed on ture populations of similar age ‘composition at the two localities separated by 250 km. ‘Theestimated biomass length) of the smallest turtles Found is five to six times les than that ofthe largest turtles found at these sites, and 16 times less than that ‘of the largest turtle carcass so far located at an eagle {feeding station. Smaller neonate turtles from 60-110 sum carapece length might be taken less frequently. ‘or consumed without leaving readily detectable rem- nants. Locality data from the 45 December 1989 prey rem- ants were combined with laity data from the 40 described in the previous report (Walker and Par ‘mente, 1990), Three measurements were taken from {he site of ech juvenile turtle record the distance to ‘the mainland (Fig. 2a the distance tothe nearest Iajor nesting colony (Peak 1 Wild Duck I Avoid {Eon the east coast and Crab. on the wesicm tp of Cape York: Fig. 20); and the regional shelf depth ‘rcluting local shallows around islands (Fig. ).Geo- {gaphicl and bathymetric characersics of juvenle Ndepresa habitat are clearly identife, although cau tion trast be exereised in inferring the absence of jveniles from other shelf areas. Many unknown or {Uncontrolled factors are involved in generation and ‘Escovery of turtle remnants including geographical ‘ariation in engle density, island density, inland size [etic search effectiveness for remnants), availabilty ofaltemative eagle prey, rainfall (efets carcass decay rteand by vegetation) terrain searching, ‘Eiealty, and available serch time. ‘The data in Fig. 2a and Fig 3 suggest that young [eter depres avold the very shallow coastal mar in. AGGIE. depresealeo ace to avoid cooeal water