Herpetologists' League

Food of Atelopus oxyrhynchus (Anura: Atelopodidae) in a Venezuelan Cloud Forest Author(s): Pedro Durant and Jim W. Dole Source: Herpetologica, Vol. 30, No. 2 (Jun., 1974), pp. 183-187 Published by: Herpetologists' League Stable URL: http://www.jstor.org/stable/3892038 . Accessed: 09/04/2011 16:36
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Acknowledgmn ts.-I

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were sighted and none were active on the partially-shaded primary sample sites. Animals aggregate for many reasons, including advantages associated with feeding, breeding, and reduction of predation. It is most likely that the U. ornatus aggregations reported here are the result of a shortage of appropriate overwintering sites. This raises several questions: What cues override territorial aggressive behavior in males allowing them to coexist in large aggregations? Perhaps cues are not necessary as this behavior may break down from hormonal changes associated with the winter nonreproductive state. Milstead (1970) observed late summer territorial behavior in U. ornatus but Martin (1973) noted that U. ornatus in Texas was nonreproductive by 17 August 1971. Since in lower deserts of Arizona these lizards are active through late October, nonreproductive associated hormonal changes could account for this territorial breakdown.

thank Drs. M. J. Fouquette, Jr. and John Alcock, Arizona State University, for criticism on the initial drafts of the manuscript.
LITERATURE CITED
MARTIN,

R. F. 1973. Reproduction in the tree lizard (Urosaurus ornatus) in central Texas: drought conditions. Herpetologica 29:27-32. MILSTEAD, W. WV.1970. Late sumer behavior of the lizards Sceloporusmerriamiand Urosaurus ornatus in the field. Herpetologica 26:343-354. VITT, L. J. 1973. Reproductive biology of the anguid lizard Gerrhonotus coeruleus prncipis. Herpetologica 29:176-184. WEINTRAUB, J. D. 1968. Winter behavior of the granite spiny lizardSceloporusorcutti Stejneger. Copeia 1968:708-712.
WORTHINGTON, R. D., AND M. D. SABATH. 1966.

aurs Winter aggregations of the lizard Uros ornatus ormatus (Baird and Girard) in Texas. Herpetologica 22:94-96. Received: 15 June 1973 Accepted: 13 August 1973

Department of Zoology, Arizona State

University, Tempe, Arizona 85281

FOOD OF ATELOPUSOXYRHYNCHUS
(ANURA: ATELOPODIDAE) IN A VENEZUELAN CLOUD FOREST
PEDRODURANT AND JINI W. DOLE
ABSTRACT: Analysis of stomach contents of 63 adult Atelopus oxyrhywhus showed that in both sexes Coleoptera accounted for approximately40% of the food biomass, while Hymenoptera (mostly ants), larval Lepido-ptera larval Diptera and Acarina constituted most of the remaining bulk. All other groups accounted for < 15%e of total weight of stomach contents. Stomachs of 9 9 usually contained almost twice as much food n aterial as did those of the smaller 8 . Frogs taken in amplexus tended to have less material in their stomachs than did unpaired animals. Of eight amplectant 8 8, six contained no identifiable food items and one contained a single beetle; average of stomach contents of these seven S S was about 25% of the average for unpaired &S. Apwveight parently 8 8, which form amplexus several months in advance of spawning, feed only infrequently if at all during this period.

NUMEROUS

studies of the food eaten by

frogs and toads of the temperate zone have been published, but our knowledge of the feeding habits of tropical anurans is more
HERPETOLOGICA 30:183-187.

scanty. Berry ( 1965, 1966), Berry and Bullock (1962) and Inger and Marx (1961) have reported on the diets of several Afncan and Asian species, but little has been

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published on the food of the anurans of tropical America. During a year-long study of the natural history and behavior of Atelopus oxyrhynchus in the cloud forest of the Venezuelan Andes we made a preliminary analysis of their stomach contents to determine the nature and quantity of food typically utilized. Because of the unusual habit in this species of initiating amplexus as much as 6 months before spawning (Dole and Durant, 1974) we were particularly interested in comparing the feeding habits of the paired and unpaired animals.
METHODS

roughly the same proportions as in the identifiable portion. All insects, arachnids and crustaceans were identified to order, and in some of the larger orders, to family. All else was identified to class. Data for both sexes and for paired and unpaired animals were treated separately.
RESULTS AND DISCUSSION

Sixty-three adult frogs (33 S ~, 30 9 9) were collected between 7 December 1970 and 23 November 1971, from the forest floor in the region surrounding our study area (see Dole and Durant, 1974 for a description of the site). Of the 63 animals 8 of each sex were in amplexus when taken, 2 on 29 March, 1 on 19 April and 5 on 7 May 1971. At capture each animal was immediately preserved in 10% Formalin. After return to the laboratory each animal was measured, its sex determined, its stomach removed and the contents washed into petri dishes. All intact, identifiable food items were then sorted by type, counted and measured. Food collections were then dried at 100 C for 24 hours and weighed to the nearest 0.01 mg. Remaining debris from the stomachs was also dried and weighed. From these data were calculated: (1) the percentage of frogs containing each type of food item (number of frogs containing any quantity of a given food item by the total number of frogs), (2) the percentage occurrence of each type of food item (number of individuals of each specific food type . the total number of all intact food items identified), and (3) the percentage weight of each item (weight of each specific food type . the total weight of all identified items). In determining this last measure the weight of unidentified debris was excluded; presumably the components of this material are in

formed the Unpaired Frogs.-Beetles largest portion of the diet of unpaired animals of both sexes. Including immature forms, Coleoptera accounted for over 40% of the identifiable biomass among the male frogs and 39% in the females (Table 1). Over 95% of the stomachs of both sexes contained adult beetles attesting to the importance of these insects in the diet. Sixty percent of the biomass of adult Coleoptera taken by males and 50% of that eaten by females was composed of snout beetles (Curculionidae); no other coleopteran family contributed more than 15% of the bulk. Hymenoptera also contributed significantly to the diet, constituting over 28% of the identifiable biomass and occurring in 90% of the stomachs of females, while in males these insects made up 20% of the weight and were found in 76% of the stomachs. Eighty-three percent of adult hymenopterans found were ants, and most of the biomass of these insects were derived from only a few frogs whose stomachs contained large numbers of individuals. One female's stomach contained 117 ants, another 79, and a third 60; these three stomachs alone accounted for 66% of all hymenopterans in the females and more than 65% of the hymenopteran biomass. Among males the largest number of ants in given stomachs were 43, 31, and 28, together accounting for 55% of all individual Hymenoptera and 70% of the hymenopteran biomass. It appears that when these frogs chance upon an ant colony they tend to feed continuously until satiated. Two other orders of insects, Diptera and Lepidoptera, contributed significantly to

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TABLE 1.-Stomach

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contents of Atelopus oxyrhynchus,by sex. (A) 25 unpaired & & and 22 unpaired 9 9 and (B) 16 pairs.
A ?9/d, _ Percent weight of item Percent of stomachs with item B ?9/Jc Percent occurrence of item Percent weight of item

Item

Percent of stomachs with item

Percent occurrence of item

INSECTA Coleoptera (adult) Coleoptera (larval) Hymenoptera (adult) Hymenoptera (larval) Diptera (adult) Diptera (larval) Lepidoptera (larval) Hemiptera Homoptera Orthoptera Trichoptera (larval) Neuroptera (larval) Thysanura Thysanoptera Collembola Embiidina Dermaptera ARACHNIDA Phalangida Acarina Araneae Pseudoscorpionida CRUSTACEA Isopoda CHILOPODA DIPLOPODA GASTROPODA

95.5/96.0 45.5/52.0 90.9/76.0 0/4.0 27.3/28.0 77.3/72.0 27.3/28.0 22.7/12.0 31.8/20.0 0/8.0 9.1/8.0 4.6/0 0/4.0 13.6/12.0 31.8/32.0 4.6/4.0 4.6/0 13.6/8.0 86.4/100.0 45.5/28.0 27.3/20.0 4.6/16.0 9.1/8.0 18.2/20.0 4.6/12.0

18.6/19.3 1.9/2.0 39.3/20.9 0/0.2 0.6/1.0 9.2/13.2 1.0/0.8 0.5/0.3 1.6/0.7 0/0.2 0.3/0.2 0.1/0 0/0.1 0.5/0.6 1.7/2.1 0.1/0.1 0.1/0 0.5/0.2 21.4/34.2 1.1/0.9 0.4/0.8 0.1/0.9 0.2/0.3 0.4/0.6 0.2/0.3

35.9/37.1 2.7/3.6 28.5/20.0 0/0.9 0.3/3.2 7.6/9.9 12.1/6.4 3.8/0.4 1.7/0.8 0/1.1 0.4/1.7 0.1/0 0/< 0.1 0.1/< 0.1 0.2/0.4 < 0.1/0.1 < 0.1/0 0.6/0.5 3.4/7.1 0.5/1.3 0.2/0.7 0.1/1.9 0.1/0.2 1.5/1.7 0.1/0.9

87.5/25.0 37.5/12.5 87.5/12.5 0/0 50.0/0 100.0/12.5 50.0/0 25.0/12.5 25.0/0 0/12.5 0/0 12.5/0 0/0 12.5/0 25.0/0 0/0 0/0 12.5/0 75.0/12.5 50.0/12.5 0/0 0/0 12.5/0 37.5/0 0/0

23.1/25.0 3.7/2.5 21.7/5.0 0/0 2.2/0 16.1/7.5 2.2/0 0.7/2.5 0.7/0 0/2.5 0/0 0.4/0 0/0 0.4/0 1.1/0 0/0 0/0 0.4/0 23.4/52.5 1.8/2.5 0/0 0/0 0.4/0 1.8/0 0/0

35.5/64.4 3.7/3.4 14.3/2.0 0/0 1.1/0 13.2/10.5 16.6/0 0.8/2.5 0.6/0 0/3.5 0/0 < 0.1/0 0/0 < 0.1/0 0.2/0 0/0 0/0 0.1/0 3.3/10.7 2.0/3.0 0/0 0/0 0.6/0 8.0/0 0/0

the diet. Only immature lepidopterans were encountered and among the dipterans, larval forms constituted the bulk of the insects taken. The small numbers of adult flies and the conspicuous absence of adult moths and butterflies is probably due to the fact that these very sluggish frogs are generally incapable of capturing them for at times they are abundant. A total of only 17 lepidopteran larvae were found, accounting for the very low percent occurrence (1% or less). In numbers of individuals, the Acarina outranked all other food items in the males (34.2% occurrence) and ranked second only

to adult hymenopterans in the females (21.4% occurrence). However, because of their small size the contribution of the mites to the biomass was only moderate, 3.4% and 7.1% of the weight of the identifiable contents in females and males respectively. Each male had at least one mite in its stomach, while 86% of the stomachs of females contained mites. The somewhat greater utilization of mites by males may be associated with the size difference between the sexes. Possibly the smaller males generally select smaller prey than the females, hence more frequently take the mites, none of which measured over 1.5 mm in length.

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2.-Mean weight of total stomach contents of paired and unpaired Atelopus oxyrhynchus.
Weight (xt + SE) in milligrams of total stomach contents % of % of total animals weight with any which identiwas iden- fiable tifiable item

3.-Analysis of variance for weights of total stomach contents of paired and unpaired Atelopus oxyrhynchusof both sexes.
Source df MS F P

paired unpaired paired unpaired

39.7 mg ? 4.5 45.4 ? 3.9 10.9 ? 3.8 28.5 ? 3.6

8 22 8 25

33.8 38.2 14.1 33.6

100 100 25 100

sex pairing interaction error

1 1 1 59

6241.8 1655.7 424.3 281.5

22.17 5.88 1.51

<.001 <.025 > .10

mals. Together these five groups accounted for 87.6% and 93.5% of the identifiable biomass in paired females and males respectively, closely approximating the comfigures (90.5% and 88.2%) for the parable Representatives of several other animal frogs. unpaired groups were also found, but none in large animals of both sexes tended to Paired numbers. All the remaining taxa together in their stomachs than material have less accounted for less than 10% of the identidid frogs (Table 2). The differunpaired fiable biomass in females and less than 12% ence significant at -Lhe2.5% is statistically in males. Of interest, however, is the pres3). On the average, stomachs level (Table ence in two males of aquatic trichopteran of only about twomales contained paired larvae, presumably indicating that they fed much those of unthirds as biomass as did in a body of water, possibly one of the in state most of it an advanced males, paired the found in lower small pools occasionally six of the of In stomachs of the digestion. areas of the forest. Since no animals were identinothing could be males eight paired taken near streams it is unlikely that they fied; a seventh frog contained a single had fed in running water. The stomachs of females on the average beetle, and only in the eighth were several contained almost twice as much material intact food items found. The stomach conas did those of males (Table 2). This dif- tents of this one animal accounted for 37.8% ference (statistically significant at the 0.1% of the total biomass of food among the level, Table 3) presumably is a reflection paired males. Eliminating this one animal of the larger size of the females (Dole and from the sample, the average weight of Durant, 1974), their greater stomach capac- the stomach contents of the remaining seven ity and their greater nutrient requirements. paired males drops to 7.7 mg or about 25% In males there was a significant positive cor- of the average for unpaired males. All unrelation between body length and total paired males contained numerous intact weight of stomach contents (r = 0.5671 food items. This very low biomass and the with 23 df; P < .01) but in females the well-digested state of the gut contents correlation lacked statistical significance among seven of the eight paired males suggests that males seldom feed after (r = 0.2868 with 20 df; P > .10). Paired Frogs.-Reduced variety in the forming amplexus. It is very likely that the stomach contents of the eight pairs of frogs male with a full stomach had only recently in amplexus reflects the smaller sample size paired. Since males in amplexus tenaciously (Table 1). All differences were among taxa and continuously grip their partners until which were only minor contributors to the spawning occurs, often several months later diet of unpaired animals. The relative con- (Dole and Durant, 1974), the emptiness of tributions of Coleoptera, Hymenoptera, their guts is not surprising. Presumably Diptera, Lepidoptera and Acarina were they subsist upon occasional arthropods similar in both paired and unpaired ani- which chance to wander across their part-

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ner's head directly in front of them, and on their fat reserves. In view of the near starvation of the males during this period the adaptive advantage of forming pairs so much in advance of spawning is difficult to see, though it may be related to the fact that males outnumber females about 3:2, thereby decreasing the probability that an unpaired male will find a mate late in the season. Females are apparently not so much affected in their feeding habits by pairing. Though paired females averaged slightly less biomass in the stomach than unpaired females, the difference was less pronounced than in males. Even after pairing the females still largely control their own movements hence have an advantage over their male partners in obtaining food. This probably accounts for the fact that paired females averaged almost 4x as much biomass in the stomach as did their partners. Presumably this difference reflects both the larger size of the females and the limitations which pairing puts on feeding by males.
thank Dr. Hernan Finol, Acknowledgments.-We Director of the Silviculture Institute for permis-

sion to use the Forestry Station, Dr. Luis Fernando Chaves V., Dr. Juan B. Castillo and Mr. Jesus Petit from the Institute of Geography for providing vehicles. Mr. Manual A. Guevara aided us on several occasions and for this we are grateful.
LITERATURE CITED BERRY,P. Y. 1965. The diet of some Singapore

anura (Amphibia). Proc. Zool. Soc. London. 144:163-174. . 1966. The food and feeding habits of the Torrentfrogs, Amolops larutensis. J. Zool. (London) 149:204-214. AND J. A. BULLOCK. 1962. The fQod of the common Malayantoad, Bufo melanostictus Schneider. Copeia 1962:736-741. DOLE, J. W., AND P. DURANT. 1974. Movements and seasonal activity of Atelopus oxyrhynchus (Anura: Atelopodidae) in a Venezuelan cloud forest. Copeia 1974:230-235. INGER, R. F., AND H. MARX. 1961. The food of amphibians. Exploration du Parc National de l'Upemba. Fascicule 64: 1-86. Received: 19 June 1973 Accepted: 7 September 1973

Facultad de Ciencias, Universidad de los Andes, Merida, Venezuela and Department of Biology, California State University, Northridge, California 91324

THE OCCURRENCE OF MULTIPLE TESTES IN THE GENUS EURYCEA (AMPHIBIA: PLETHODONTIDAE)

DAVID M. SEVER
ABSTRACT: Multipletestes in urodelesconsistof a seriesof enlargements whichwhenfully developed aremorphologically and functionally to each other or to a simple testis. Such testes are known similar within the tribe Hemidactyliini E. nana,andE. neotenes.Studies for only Eurycealucifuga, on the morphogenesis of multipletestesare neededto settle conflictingreportsconcerning their formation and to ascertain and regression definitelythe time intervalbetweenformation of successive testicular lobes.

A MULTIPLE testis is made up of a series of enlarged lobes that are linearly arranged and separated by constricted areas that may be of greater length than the lobular enlargements (Humphrey, 1922). Multiple testes have been described in Desmognathus (Kingsbury, 1902; Humphrey, 1921, 1922;
HERPETOLOGICA 30:187-193. June

and Burger, 1937), Salamwndra (Kingsbury, 1902; Nussbaum, 1906; Champy, 1913; Humphrey, 1922; Baker, 1965), Notophthalmus (Humphrey, 1922, 1926; Obreshkove, 1924; Adams, 1940; Baker, 1965), Taricha (Humphrey, 1922; McCurdy, 1931; Miller and Robbins, 1954; Baker, 1965), Leurog-

1974