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diff --git a/33574.txt b/33574.txt new file mode 100644 index 0000000..273a188 --- /dev/null +++ b/33574.txt @@ -0,0 +1,1663 @@ +The Project Gutenberg EBook of A Field Study of the Kansas Ant-Eating +Frog, Gastrophryne olivacea, by Henry S. Fitch + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: A Field Study of the Kansas Ant-Eating Frog, Gastrophryne olivacea + +Author: Henry S. Fitch + +Release Date: August 29, 2010 [EBook #33574] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK KANSAS ANT-EATING FROG *** + + + + +Produced by Simon Gardner, Chris Curnow, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + +Transcriber's Notes + +In this Plain Text version of the book, italic typeface is represented +with _underscores_, and small capital typeface is represented in UPPER +case. + + [=e] represents a macron (horizontal line) over an e. + [~n] represents n-tilde. + [Female] represents the symbol for female. + [Male] represents the symbol for male. + +A small number of inconsistencies and typographical errors have been +changed in the text. These are listed at the end of this book. + +The Title page and Verso are in error in stating that the pages run 275 +to 306. This should read 276-307. + + * * * * * + + + UNIVERSITY OF KANSAS PUBLICATIONS + + MUSEUM OF NATURAL HISTORY + + Volume 8, No. 4, pp. 275-306, 9 figs. in text + + February 10, 1956 + + + A Field Study + of the Kansas Ant-Eating Frog, + Gastrophryne olivacea + + + BY + + HENRY S. FITCH + + + UNIVERSITY OF KANSAS + LAWRENCE + 1956 + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, A. Byron Leonard, + Robert W. Wilson + + + + Volume 8, No. 4, pp. 275-306, 9 figs. in text + Published February 10, 1956 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + FERD VOILAND. JR., STATE PRINTER + TOPEKA, KANSAS + 1956 + + 25-7819 + + + + +A FIELD STUDY OF THE KANSAS ANT-EATING FROG, GASTROPHRYNE OLIVACEA + +By + +Henry S. Fitch + + +INTRODUCTION + +The ant-eating frog is one of the smallest species of vertebrates on the +University of Kansas Natural History Reservation, but individually it is +one of the most numerous. The species is important in the over-all +ecology; its biomass often exceeds that of larger species of +vertebrates. Because of secretive and subterranean habits, however, its +abundance and effects on community associates are largely obscured. + +The Reservation, where my field study was made, is the most northeastern +section in Douglas County, Kansas, and is approximately 5-1/2 miles north +and 2-1/2 miles east of the University campus at Lawrence. The locality +represents one of the northernmost occurrences of the species, genus, +and family. The family Microhylidae is a large one, and most of its +representatives are specialized for a subterranean existence and a diet +of termites or ants. The many subfamilies of microhylids all have +distributions centering in the regions bordering the Indian Ocean, from +South Africa and Madagascar to the East Indies, New Guinea, and +Australia (Parker, 1934). Only one subfamily, the Microhylinae, is +represented in the New World, where it has some 17 genera (de Carvalho, +1954) nearly all of which are tropical. _G. olivacea_, extending north +into extreme southern Nebraska (Loomis, 1945: 211), ranges farther north +than any other American species. In the Old World only _Kaloula +borealis_ has a comparable northward distribution. Occurring in the +vicinity of Peiping (Pope, 1931: 587), it reaches approximately the same +latitude as does _Gastrophryne_ in Nebraska. The great majority of +microhylid genera and species are confined to the tropics. + +Nearly all ant-eating frogs seen on the Reservation have been caught and +examined and individually marked. By November 1, 1954, 1215 individuals +had been recorded with a total of 1472 captures. In the summer of 1950, +Richard Freiburg studied this frog on the Reservation and his findings +(1951) led to a better understanding of its natural history. The +numbers of frogs studied by him however, were relatively small and the +field work was limited to the one summer. The data now at hand, +representing six consecutive years, 1949 through 1954, serve to +supplement those obtained by Freiburg, corroborating and extending his +conclusions in most instances, and also indicating that certain of his +tentative conclusions need to be revised. + +While the present report was in preparation, Anderson (1954) published +an excellent account of the ecology of the eastern species _G. +carolinensis_ in southern Louisiana. Anderson's findings concerning this +closely related species in a much different environment have been +especially valuable as a basis for comparison. The two species are +basically similar in their habits and ecology but many minor differences +are indicated. Some of these differences result from the differing +environments where Anderson's study and my own were made and others +certainly result from innate genetic differences between the species. + +The frog with which this report is concerned is the _Microhyla +carolinensis olivacea_ of the check list (Schmidt, 1953: 77) and recent +authors. De Carvalho (1954: 12) resurrected the generic name, +_Gastrophryne_, for the American species formerly included in +_Microhyla_, and presented seemingly valid morphological evidence for +this plausible generic separation. + +_G. olivacea_ is obviously closely related to _G. carolinensis_; the +differences are not greater than those to be expected between well +marked subspecies. Nevertheless, in eastern Oklahoma and eastern Texas, +where the ranges meet, the two kinds have been found to maintain their +distinctness, differing in coloration, behavior, calls, and time of +breeding. Hecht and Matalas (1946: 2) found seeming intergrades from the +area of overlapping in eastern Texas, but some specimens from this same +area were typical of each form. Their study was limited to preserved +material, in which some characters probably were obscured. More field +work throughout the zone of contact is needed. The evidence of +intergradation obtained so far seems to be somewhat equivocal. + +Besides _G. olivacea_ and typical _G. carolinensis_ there are several +named forms in the genus, including some of doubtful status. The name +_mazatlanensis_ has been applied to a southwestern population, which +seems to be a well marked subspecies of _olivacea_, but as yet +_mazatlanensis_ has been collected at few localities and the evidence of +intergradation is meager. The names _areolata_ and _texensis_ have been +applied to populations in Texas. Hecht and Matalas (1946: 3) consider +_areolata_ to be a synonym of _olivacea_, applied to a population +showing intergradation with _carolinensis_, but Wright and Wright (1949: +568) consider _areolata_ to be a distinct subspecies. _G. texensis_ +generally has been considered to be a synonym of _olivacea_. Other +species of the genus include the tropical _G. usta_, _G. elegans_ and +_G. pictiventris_. + +Of the vernacular names hitherto applied to _G. olivacea_ none seems +appropriate; I propose to call the species the Kansas ant-eating frog +because of its range extending over most of the state, and because of +its specialized food habits. The type locality, originally stated to be +"Kansas and Nebraska" (Hallowell, 1856: 252) has been restricted to Fort +Riley, Kansas (Smith and Taylor, 1950: 358). Members of the genus have +most often been referred to as toads rather than frogs because of their +more toadlike appearance and habits. However, this family belongs to the +firmisternial or froglike division of the Salientia and the terms "frog" +and "toad," originally applied to _Rana_ and _Bufo_ respectively, have +been extended to include assemblages of related genera or families. +Members of the genus and family usually have been called +"narrow-mouthed" toads from the old generic name _Engystoma_, a synonym +of _Gastrophryne_. _G. olivacea_ usually has been referred to as the +Texas narrow-mouthed toad, or western narrow-mouthed toad. The latter +name is inappropriate because the geographic range is between that of a +more western representative (_mazatlanensis_) and a more eastern one +(_carolinensis_). The names _texensis_, _areolata_ and _carolinensis_ +have all been applied to populations in Texas, and it is questionable +whether typical _olivacea_ even extends into Texas. + + +HABITAT + +In the northeastern part of Kansas at least, rocky slopes in open woods +seem to provide optimum habitat conditions. This type of habitat has +been described by several earlier workers in this same area, Dice (1923: +46), Smith (1934: 503) and Freiburg (1951: 375). Smith (1950: 113) +stated that in Kansas this frog is found in wooded areas, and that rocks +are the usual cover, but he mentioned that outside of Kansas it is often +found in mesquite flats that are devoid of rocks. Freiburg's field work +was done almost entirely on the Reservation and was concentrated in +"Skink Woods" and vicinity, where much of my own field work, both before +and afterward, was concentrated. On the Reservation and in nearby +counties of Kansas, the habitat preferences of the ant-eating frog and +the five-lined skink largely coincide. In an account of the five-lined +skink on the Reservation, I have described several study areas in some +detail (Fitch, 1954: 37-41). It was on these same study areas (Quarry, +Skink Woods, Rat Woods) that most of the frogs were obtained. + +Although _G. olivacea_ thrives in an open-woodland habitat in this part +of its range, it seems to be essentially a grassland species, and it +occurs throughout approximately the southern half of the Great Plains +region. Bragg (1943: 76) emphasized that in Oklahoma it is widely +distributed over the state, occupying a variety of habitats, with little +ecological restriction. Bragg noted, however, that the species is +rarely, if ever, found on extensive river flood plains. On various +occasions I have heard _Gastrophryne_ choruses in a slough two miles +south of the Reservation. This slough is in the Kaw River flood plain +and is two miles from the bluffs where the habitat of rocky wooded +slopes begins that has been considered typical of the species in +northeastern Kansas. It seems that the frogs using this slough are not +drawn from the populations living on the bluffs as Mud Creek, a Kaw +River tributary, intervenes. The creek channel at times of heavy +rainfall, carries a torrent of swirling water which might present a +barrier to migrating frogs as they are not strong swimmers. The frogs +could easily find suitable breeding places much nearer to the bluffs. +Those using the slough are almost certainly permanent inhabitants of the +river flood plain. The area in the neighborhood of the slough, where the +frogs probably live, include fields of alfalfa and other cultivated +crops, weedy fallow fields, and the marshy margins of the slough. In +these situations burrows of rodents, notably those of the pocket gopher +(_Geomys bursarius_), would provide subterranean shelter for the frogs, +which are not efficient diggers. + +The frogs may live in many situations such as this where they have been +overlooked. In the absence of flat rocks providing hiding places at the +soil surface, the frogs would rarely be found by a collector. The volume +and carrying quality of the voice are much less than in other common +anurans. Large breeding choruses might be overlooked unless the observer +happened to come within a few yards of them. Most of the recorded +habitats and localities of occurrence may be those where the frog +happens to be most in evidence to human observers, rather than those +that are limiting to it or even typical of it. + +On September 20, 1954, after heavy rains, juveniles dispersing from +breeding ponds were in a wide variety of situations, including most of +the habitat types represented on the Reservation. Along a small dry +gully in an eroded field formerly cultivated, and reverted to tall +grass prairie (big bluestem, little bluestem, switch grass, Indian +grass), the frogs were numerous. Many of them were flushed by my +footsteps from cracks in the soil along the gully banks. In reaching +this area the frogs had moved up a wooded slope from the pond, crossed +the limestone outcrop area at the hilltop edge, and wandered away from +the woods and rocks, out into the prairie habitat. In this prairie +habitat there were no rocks providing hiding places at the soil surface, +but burrows of the vole (_Microtus ochrogaster_) and other small rodents +provided an abundance of subterranean shelter. In the summer of 1955 the +frogs were seen frequently in this same area, especially when the soil +was wet from recent rain. When the surface of the soil was dry, none +could be found and presumably all stayed in deep cracks and burrows. + +Anderson (1954: 17) indicated that _G. carolinensis_ in Louisiana +likewise occurs in diverse habitats, being sufficiently adaptable to +satisfy its basic requirements in various ways. + + +BEHAVIOR + +Ordinarily the ant-eating frog stays beneath the soil surface, in cracks +or holes or beneath rocks. Probably it obtains its food in such +situations, and rarely wanders on the surface. The occasional +individuals found moving about above ground are in most instances +flushed from their shelters by the vibrations of the observer's +footsteps. On numerous occasions I have noticed individuals, startled by +nearby footfalls, dart from cracks or under rocks and scuttle away in +search of other shelter. Such behavior suggests that digging predators +may be important natural enemies. The gait is a combination of running +and short hops that are usually only an inch or two in length. The flat +pointed head seems to be in contact with the ground or very near to it +as the animal moves about rapidly and erratically. The frog has a +proclivity for squeezing into holes and cracks, or beneath objects on +the ground. The burst of activity by one that is startled lasts for only +a few seconds. Then the frog stops abruptly, usually concealed wholly or +in part by some object. Having stopped it tends to rely on concealment +for protection and may allow close approach before it flushes again. + +Less frequently, undisturbed individuals have been seen wandering on the +soil surface. Such wandering occurs chiefly at night. Diurnal wandering +may occur in relatively cool weather when night temperatures are too low +for the frogs to be active. Wandering above ground is limited to times +when the soil and vegetation are wet, mainly during heavy rains and +immediately afterward. + +Pitfalls made from gallon cans buried in the ground with tops open and +flush with the soil surface were installed in 1949 in several places +along hilltop rock outcrops where the frogs were abundant. The number of +frogs caught from day to day under varying weather-conditions provided +evidence as to the factors controlling surface activity. After nights of +unusually heavy rainfall, a dozen frogs, or even several dozen, might be +found in each of the more productive pitfalls. A few more might be +caught on the following night, and occasional stragglers as long as the +soil remained damp with heavy dew. Activity is greatest on hot summer +nights. Below 20 deg. C. there is little surface activity but individuals +that had body temperatures as low as 16 deg. C. have been found moving +about. + +Frogs uncovered in their hiding places beneath flat rocks often remained +motionless depending on concealment for protection, but if further +disturbed, they made off with the running and hopping gait already +described. Although they were not swift, they were elusive because of +their sudden changes of direction and the ease with which they found +shelter. When actually grasped, a frog would struggle only momentarily, +then would become limp with its legs extended. The viscous dermal +secretions copiously produced by a frog being handled made the animal so +slippery that after a few seconds it might slide from the captor's +grasp, and always was quick to escape when such an opportunity was +presented. + + +TEMPERATURE RELATIONSHIPS + +Ant-eating frogs are active over a temperature range of at least 16 deg. +C. to 37.6 deg. C. They tolerate high temperatures that would be lethal +to many other kinds of amphibians, but are more sensitive to low +temperatures than any of the other local species, and as a result their +seasonal schedule resembles that of the larger lizards and snakes more +than those of other local amphibians. The latter become active earlier +in the spring. + +Earliest recorded dates when the frogs were found active in the course +of the present study from 1950 to 1955 were in April every year; the +20th, 25th, 24th, 2nd, 25th, and 21st. Latest dates when the frogs were +found in the six years of the study were: October 22, 1949; October 13, +1950; October 7, 1951; August 24, 1952; August 18, 1953; and October 27, +1954 (excluding two late stragglers caught in a pitfall on December 5). +Severe drought caused unseasonably early retirement in 1952 and 1953. + +Body temperatures of the frogs were taken with a small mercury +thermometer of the type described by Bogert (1949: 197); the bulb was +used to force open the mouth and was thrust down the gullet into the +stomach. To prevent conduction of heat from the hand, the frog was held +down through several layers of cloth, at the spot where it was +discovered, until the temperature reading could be made. This required +approximately five seconds. + +[Illustration: FIG. 1. Temperatures of ant-eating frogs grouped in +one-degree intervals; upper figure is of frogs found active in the open, +and lower is of those found under shelter. The frogs are active over a +temperature range of more than 20 degrees, and show no clear cut +preference within this range.] + +Most of the 79 frogs of which temperatures were measured, were found +under shelter, chiefly beneath flat rocks. The rocks most utilized were +in open situations, exposed to sunshine. Most of the frogs were in +contact with the warmed undersurfaces of such rocks. Forty-three of the +frogs, approximately 54.5 percent, were in the eight-degree range +between 24 deg. and 31 deg. C. Probably the preferred temperatures lie +within this range. The highest body temperature recorded, 37.6 deg. C., +was in a frog which "froze" and remained motionless in the sunshine for +half a minute after the rock sheltering it was overturned. Probably its +temperature was several degrees lower while it was sheltered by the +rock. Other unusually high temperatures were recorded in newly +metamorphosed frogs found hiding in piles of decaying vegetation near +the edge of the pond, on hot afternoons of late August. Temperatures +ranged from 17.0 deg. to 30.7 deg. in frogs that were found actually +moving about. Several with relatively low temperatures, 22 deg. to 17 +deg., were juveniles travelling in rain or mist on cool days. These +frogs, having relatively low temperature, were sluggish in their +movements, as compared with individuals at the upper end of the +temperature range. + +[Illustration: FIG. 2. Body temperatures and nearby air temperatures for +frogs found under natural conditions. Dots represent frogs found under +shelter; circles represent those found in the open.] + +After the first frost each year the frogs usually could not be found, +either in the open or in their usual hiding places beneath rocks. They +probably had retired to deep subterranean hibernation sites. The only +exception was in 1954, when two immature frogs were found together in a +pitfall on the morning of December 5 after a rain of .55 inches ending +many weeks of drought. Air temperature had been little above 10 deg. C. +that night, but had often been below freezing in the preceding five +weeks. + +Reactions of these same two individuals to low temperatures were tested +in the laboratory. At a body temperature of 11 deg. C. they were +extremely sluggish. They were capable of slow, waddling movements, but +were reluctant to move and tended to crouch motionless. Even when they +were prodded, they usually did not move away, but merely flinched +slightly. At 6 deg. C. they were even more sluggish, and seemed +incapable of locomotion, as they could not be induced to hop or walk by +prodding with a fine wire. When placed upside down on a flat surface, +they could turn over, but did so slowly, sometimes only after a minute +or more had elapsed. Respiratory throat movements numbered 46 and 60 per +minute. + + +BREEDING + +Many observers have noted that breeding activity is initiated by heavy +rains in summer. In my experience precipitation of at least two inches +within a few days is necessary to bring forth large breeding choruses. +With smaller amounts of precipitation only stragglers or small +aggregations are present at the breeding ponds. Tanner (1950: 48) stated +that in three years of observation, near Lawrence, Kansas, the first +storms to bring large numbers of males to the breeding ponds occurred on +June 20, 1947, June 18, 1948, and May 1, 1949. + +In 1954 the frogs were recorded first on April 25, but these were under +massive boulders, and were still semi-torpid. Frogs were found fully +active, in numbers, under small flat rocks on May 7. They were found +frequently thereafter. On the afternoon of May 13, the third consecutive +day with temperature slightly above 21 deg. C., low croaking of a frog +was heard among rocks at an old abandoned quarry. Throughout the +remainder of May, calling was heard frequently at the quarry on warm, +sunny afternoons. Often several were calling within an area of a few +square yards, answering each other and maintaining a regular sequence. +In the last week of May rains were frequent, and the precipitation +totalled 2.09 inches. On June 1 and 2 also, there were heavy rains +totalling 2.26 inches. On the evening of June 2 many frogs were calling +at a pond 1/2 mile south of the Reservation, and one was heard at the +pond on the Reservation. By the evening of June 4, dozens were calling +in shallow water along the edge of this pond in dense _Polygonum_ and +other weeds. There was sporadic calling even in daylight and there was a +great chorus each evening for the next few days, but its volume rapidly +diminished. + +In mid-June a system of drift fences and funnel traps was installed 200 +yards west of the pond in the dry bottom of an old diversion ditch +leading from the pond. The ditch constituted the boundary between +bottomland pasture and a wooded slope, and therefore was a natural +travelway. The object of the installation was to intercept and catch +small animals travelling along the ditch bottom. The drift fence was +W-shaped, with a funnel trap at the apex of each cone so that the +animals travelling in either direction would be caught. The numbers of +frogs caught from time to time during the summer provided information as +to their responses to weather in migrating to the pond. + +TABLE 1. NUMBERS OF FROGS CAUGHT WITHIN TWO DAYS AFTER RAIN IN FUNNEL +TRAPS IN 1954, FROM MID-JUNE, TO THE TIME OF FIRST FROST. + + + Date Precipitation No. of + in inches caught frogs + July 1 2.02 8 + July 10 .11 none + July 16 1.26 none + July 20-21 .94 3 + July 24 .38 2 + July 28 .29 none + August 1-2 3.22 31 + August 6-7-8 2.43 none + August 12 .28 none + August 16 .29 none + August 19-22 .70 none + August 27-28 1.05 none + September 9 .50 none + September 29-30 .38 none + October 4 .74 none + October 12-14 3.51 none + +From the positions of the traps and drift fences, it was obvious that +all of the frogs that were caught were travelling toward the pond. +Capture of an equal number moving away from the pond a few days +afterward might have been expected but none at all was caught while +making a return trip. Therefore it seems that the frogs returned by a +different route to their home ranges after breeding. Of necessity they +make the return trip under conditions drier than those that prevail on +the pondward trip, which is usually made in a downpour. Probably the +return travel is slower, more leisurely, and with more tendency to keep +to sheltered situations. + +The call is a bleat, resembling that of a sheep, but higher, of lesser +volume, and is not unlike the loud rattling buzz of an angry bee. The +call is usually of three to four seconds duration, with an interval +several times as long. Calling males were floating, almost upright, in +the water within a few yards of shore, where there was dense vegetation. +The throat pouch when fully expanded is several times as large as the +entire head. When a person approached to within a few yards of frogs +they usually stopped calling, submerged, and swam to a place of +concealment. + +Having heard the call of typical _G. carolinensis_ in Louisiana, I have +the impression that it is a little shorter, more sheeplike, and less +insectlike than that of _G. olivacea_. The call of _Gastrophryne_ is of +such peculiar quality that it is difficult to describe. Different +observers have described it in different terms. Stebbins (1951: 391) has +described the call in greatest detail, and also has quoted from the +descriptions of it previously published. These descriptions include the +following: "high, shrill buzz"; "buzz, harsh and metallic"; "like an +electric buzzer"; "like bees at close range but more like sheep at a +distance"; "bleating baa"; "shrill, long-drawn quaw quaw"; "whistled +wh[=e][=e] followed by a bleat." + +Stebbins observed breeding choruses (_mazatlanensis_) at Pe[~n]a Blanca +Springs, Arizona, and stated that sometimes three or four called more or +less together, but that they seldom started simultaneously. Occasionally +many voices would be heard in unison followed by an interval of silence, +but this performance was erratic. At the pond on the Reservation I noted +this same tendency many times. After a lull the chorus would begin with +a few sporadic croaks, then four or five or even more frogs would be +calling simultaneously from an area of a few square yards. Anderson +(_op. cit._: 34) found that in small groups of calling _G. carolinensis_ +there was a distinct tendency to maintain a definite pattern in the +sequence of the calls. One "dominant" individual would initiate a series +of calls, and others each in turn would take up the chorus. + +Pairing takes place soon after the breeding aggregations are formed. On +the night of June 4, 1954, a clasping pair was captured and kept in the +laboratory in a large jar of water. This pair did not separate, and +spawning occurred between noon and 1:30 P. M. on June 5. When the newly +laid eggs were discovered at 1:30 P. M. most of them were in a surface +film. Some were attached to submerged leaves and a few rested on the +bottom. The pair was still joined, but the male was actually clasping +only part of the time, and as the frogs moved about in the water, it +became evident that they were adhering to each other by the areas of +skin contact, which were glued together by their dermal secretion. They +were unable to separate immediately, even when they struggled to do so. +They were observed for approximately 15 minutes before separation +occurred, and during this time they were moving about actively. As they +separated, the area of adhesion was discernible on the back of the +female. It was U-shaped, following the ridges of the ilia and the +sacrum. + +On August 2, 1954, after a rain of 3.22 inches, the previously mentioned +funnel trap in the ditch had caught 31 ant-eating frogs. Water had +collected to a depth of several inches in the depression where the trap +was situated. A dozen of the trapped frogs were clasping pairs. These +frogs struggled vigorously as they were removed from the traps, handled +and marked. As a result most of the clasping males were separated from +the females. In handling those of each pair I noticed that they were +glued together by dermal secretions, as were those of the pair observed +on June 5. The areas of adhesion were of similar shape and location in +the different pairs, and included the U-shaped ridge of the female's +back and the male's belly, and the inner surfaces of the male's forelegs +with the corresponding surfaces of the female's sides where the male +clasped. + +This adhesion of the members of a pair during mating may be a normal +occurrence. The copious secretion of the dermal glands is of especially +glutinous quality in _Gastrophryne_. The adhesion of members of a pair +may have survival value. These small frogs are especially shy, and in +the breeding ponds they respond to any disturbance with vigorous +attempts to escape and hide. Under such circumstances the adhesion may +prevent separation. Also, it may serve to prevent displacement of a +clasping male by a rival. Anderson (_op. cit._) who observed many +details of the mating behavior of _G. carolinensis_, both in the +laboratory and under natural conditions, mentioned no such adhesion +between members of a pair. + +Anderson (_op. cit._: 31) discussed the possibility that reproductive +isolation might arise in sympatric populations, such as those of _G. +carolinensis_ in southern Louisiana, through inherent differences in +time of spawning. However, in _G. olivacea_ at least, such isolation +would be prevented by individual males returning to breed at different +times in the same season. Furthermore, individual differences in choice +of breeding time probably result from environmental factors rather than +genetic factors in most instances. In _G. olivacea_ in Kansas, time of +breeding is controlled by the distribution of heavy rainfall creating +favorable conditions. Onset of the breeding season may be hastened or +delayed, or an entire year may be missed because of summer drought. If +favorable heavy rains are well distributed throughout the summer, frogs +of age classes that are not yet sexually mature in the early part of +the breeding season, may comprise the bulk of the breeding population in +late summer. + + +DEVELOPMENT OF EGGS AND LARVAE + +Eggs laid on June 5 by the pair kept in the laboratory were hatching on +June 7, on the average approximately 48 hours from the time of laying. +By June 8 all the eggs had hatched and the tadpoles were active. On +August 28 and 29 thousands of newly metamorphosed young were in evidence +on wet soil at the pond margin; in some the head still was tadpolelike +and they had a vestige of the tail stump. These young were remarkably +uniform in size, 15 to 16 mm. (the smallest one found was 14-1/2 mm.) +and almost all of them had originated from eggs laid after heavy +precipitation, totalling 3.22 inches, in the first 36 hours of August. +Allowing one day for adults to reach the pond and spawn, and two days +more for eggs to hatch, the tadpole stage must have lasted approximately +24 days in this crop of young. + +Wright and Wright (1949: 582) stated that the tadpoles metamorphosed +after 30 to 50 days, and that the newly metamorphosed frogs are 10 to 12 +mm. in length. Length of time required for larval development probably +varies a great deal depending on the interaction of several factors such +as temperature and food supply. + + +GROWTH + +Little has been recorded concerning the growth rate of _Gastrophryne_ or +the time required for it to attain sexual maturity. Wright (1932) found +that _G. carolinensis_ in the Okefinokee Swamp region has a mean +metamorphosing-size of 10.8 mm. Young thought to be those recently +emerged from their first hibernation were those in the size group 15.0 +to 20.0 mm., while the frogs in the 20 to 27 mm. size class and those in +the 27 to 36 mm. class were interpreted as representing two successively +older annual age classes. Anderson (1954: 41) thought he could recognize +four successive annual age classes in the same species in southern +Louisiana. He found that sexual maturity is attained at a length of 21 +to 24 mm. in frogs which he believed to be late in the second year of +life. + +Allowing for size differences between the two species, Wright's and +Anderson's conclusions regarding growth in _G. carolinensis_, on the +basis of size groups, are largely substantiated by my own data on the +growth of marked individuals of _G. olivacea_ living under natural +conditions in Kansas. + +In 1954, an opportunity to investigate the early growth was afforded by +unusually favorable circumstances. The population of frogs that emerged +from hibernation in the late spring of 1954 included few, if any, that +were below adult size; drought had prevented successful breeding in 1952 +and 1953. Heavy rains in the first week of June, 1954, and again in the +first week of August, resulted in the production of two successive crops +of young so widely spaced that they were easily distinguishable. Some +young may have been hatched after other minor rains, but certainly these +were relatively few. Young from the eggs laid in the first week of +August were metamorphosing during the last week of August. Growth in the +frogs of this group can be shown by the average size and the size range +of the successive samples collected. + +TABLE 2. GROWTH IN FROGS METAMORPHOSED IN THE LAST WEEK OF AUGUST, 1954. + + =========================================================== + |Number in| Mean size |Size range + Time of sample | sample | in mm. | in mm. + --------------------+---------+----------------+----------- + August 27 to 31 | 27 | 15.55 +/- .079 | 15 to 17 + --------------------+---------+----------------+----------- + September 11 | 114 | 17.2 +/- .033 | 14 to 20 + --------------------+---------+----------------+----------- + September 15 to 22 | 12 | 18.7 +/- .090 | 16 to 20 + --------------------+---------+----------------+----------- + September 27 to 30 | 37 | 19.3 +/- .055 | 17 to 21.5 + --------------------+---------+----------------+----------- + October 1 to 7 | 62 | 20.8 +/- .072 | 17 to 24 + --------------------+---------+----------------+----------- + October 12 to 17 | 49 | 22.3 +/- .092 | 18 to 24 + =========================================================== + +By mid-October, six weeks after metamorphosis, these frogs had increased +in over-all length by approximately 50 percent. Having grown a little +more than 1 mm. per week on the average, they were approximately +intermediate in size between small adults and newly metamorphosed young. + +The frogs hatched in June were present in relatively small numbers +compared with those hatched in August, and were not observed +metamorphosing. In late August a sample of 33 judged to belong to the +June brood averaged 26.2 (22-28) mm. long. A sample of 39 from the first +week of October averaged 28.1 (24.5-32) mm. Frogs of this group thus +were approaching small adult size late in their first growing season. +Such individuals possibly breed in the summer following their first +hibernation, when they are a year old or a little more. Because +recaptured frogs were not sacrificed to determine the state of their +gonads, the minimum time required to attain sexual maturity was not +definitely determined. The available evidence indicates that sexual +maturity is most often attained late in the second year of life, at an +age of approximately two years. The darkened and distensible throat +pouch of the adult male probably is the best available indicator of +sexual maturity. + +[Illustration: FIG. 3. Growth shown by successive samples of young +ant-eating frogs of two size groups in late summer and early fall of +1954. For each sample the mean, standard deviation, and range are shown. +Lower series are those metamorphosed in late August, and upper series +are those metamorphosed in late June.] + +[Illustration: FIG. 4. Rapid growth of a young female caught in June, +July, and August, 1949. Presumably this individual metamorphosed late in +the summer of 1948, and at the age of approximately one year it was near +small adult size.] + +Frogs that metamorphose in late summer have little time to grow before +hibernating, and still are small when they emerge in spring. The +smallest one found was 19 mm. long (May 19, 1951), and in each year +except 1954 many such young were found that were less than 25 mm. in +length in May or early June. None of the frogs marked at or near +metamorphosing size has been recaptured, but the trend of early growth +is well shown by Table 2 and Fig. 3. However, many juveniles that were +captured and marked within a few weeks of metamorphosis were recaptured +as adults. The selected individuals in Table 3 are considered typical of +growth from "half-grown" to small adult size. Growth in many other +individuals is shown in Figs. 6 and 7. + +TABLE 3. GROWTH IN FROGS MARKED AS YOUNG AND RECAPTURED AS SMALL ADULTS. + + ============================================================== + Individual | Dates | Length | Probable time + and sex | of capture | in mm. |of metamorphosis + -----------------+-----------------+---------+---------------- + No. 1 [Female] | August 28, 1951 | 21.5 |Mid-July, 1951 + | May 5, 1952 | 23 | + | July 3, 1952 | 32 | + | August 31, 1952 | 33 | + -----------------+-----------------+---------+---------------- + No. 2 [Female] | June 8, 1950 | 25 |Late July, 1949 + | May 24, 1951 | 31 | + | July 30, 1951 | 34 | + | June 24, 1952 | 35 | + -----------------+-----------------+---------+---------------- + No. 3 [Male] | August 31, 1951 | 24 |Late June, 1951 + | May 23, 1953 | 32 | + ============================================================== + +[Illustration: FIG. 5. Ant-eating frogs, a little less than twice +natural size, adult and newly metamorphosed young, showing differences +in size and coloration. The young is darker and has a leaflike middorsal +mark which fades as growth proceeds.] + +The trend of growth after attainment of minimum adult size is also well +shown by the records of marked individuals recaptured. Many of these +were marked while they were still small so that their approximate ages +are known. For those recaptured in their second year, after one +hibernation, length averaged 30.92 mm. Some of this group were young +metamorphosed late the preceding summer and still far short of adult +size (as small as 23 mm.) when recaptured. Others were relatively large, +up to 33 mm. A group of 22 recaptured frogs known to be in their third +year averaged 33.3 mm. (males 31.9, females 35.3, excluding four +individuals of undetermined sex). Fifteen other recaptured frogs were +known to be in their fourth year at least, and some probably were older, +as they were already large adults when first examined. These 15 averaged +36.6 mm. (males 34.7, females 37.9 mm.). Size was similar in a sample of +58 individuals intercepted en route to the breeding pond in heavy rains +of June and August, 1954. The 38 males in this sample ranged in size +from 30 mm. to 38 mm., averaging 34.5. The 20 females ranged from 34 mm. +to 40 mm., averaging 37.65. The large average and maximum size in this +sample of a breeding population may be typical after periods of drought +years have prevented successful reproduction. Summer drought in 1952 and +1953 prevented breeding in those years, or, at least, it drastically +reduced the numbers of young produced. One-year-old and two-year-old +frogs may not have been represented at all in the sample of 58. +Three-year-old frogs presumably made up a substantial part of the +sample, since 1951 was a year of successful breeding. + +[Illustration: FIG. 6. Growth in a group of frogs, each marked while +still short of adult size and mostly recaptured after lapse of one or +more hibernation periods. Each line connects records of an individual +frog.] + +Differences in size between species and geographic variation in size in +_Gastrophryne_ have been given little attention by herpetologists, but +if understood, would help to clarify relationships. Hecht and Matalas +stated in their revision (1946: 5) that size is of no importance as a +taxonomic character, as typical _carolinensis_, _olivacea_, and +_mazatlanensis_ all averaged approximately the same--26 to 28 +mm.--females slightly larger than males. However, they arbitrarily +classed as adults all individuals 22.5 mm. in length or larger, having +found individuals this small that showed the darkened and distensible +throat pouches characteristic of adult males. From the trend of my own +measurements of _G. olivacea_ in northeastern Kansas, I conclude that +either many immature individuals were included in their samples, or that +the populations sampled included some with individuals that were +remarkably small as adults. + +[Illustration: FIG. 7. Growth in another group of frogs that were marked +as young or small adults and recaptured after intervals of more than a +year. Frogs of this group were, on the average, larger than the +individuals shown in Fig. 6, and they made less rapid growth.] + +The population which I studied may be considered typical of _G. +olivacea_. They averaged large, including individuals up to 42 mm. in +length, well above the maximum sizes for any reported in the literature. +At metamorphosis these _olivacea_ are of approximately 50 percent +greater length than _G. carolinensis_ as reported by Wright and Wright +(1949: 573) and Anderson (1954: 41). Yet Blair (1950: 152) observed that +in eastern Oklahoma, where the ranges of _olivacea_ and _carolinensis_ +overlap, the latter is larger. On the basis of field and laboratory +observations he tentatively concluded that one of the main barriers to +interbreeding was the reluctance of the males of _carolinensis_ to clasp +the smaller females of _olivacea_. + +That size differs in different populations, and is still poorly +understood, is illustrated by the following discrepant figures from +various authors. + +TABLE 4. SIZE RANGE OF ADULTS IN VARIOUS POPULATIONS OF GASTROPHRYNE. + + ===============+=======================+=================+============= + Species or | Geographic population | Authority |Size range of + subspecies | sampled | |adults in mm. + ---------------+-----------------------+-----------------+------------- + | | | + _olivacea_ |Douglas Co., Kansas |present study | 31 to 42 + | | | + _olivacea_ |entire range |Wright and Wright| 19 to 38 + | | (1949) | + | | | + _carolinensis_ |entire range |Wright and Wright| 20 to 36 + | | (1949) | + | | | + _carolinensis_ |southern Louisiana |Anderson | 22 to 35 + | | (1954) | + | | | + _areolata_ |southeastern Texas |Wright and Wright| 23 to 29 + | | (1949) | + | | | + _mazatlanensis_|Arizona and New Mexico |Wright and Wright| 22 to 30 + | | (1949) | + | | | + _mazatlanensis_|Santa Cruz Co., Arizona|Stebbins | 25.2 to 31.5 + | | (1951) | + ---------------+-----------------------+-----------------+------------- + + +COLOR AND PATTERN + +The color pattern changes in the course of development, and the shade of +color changes in response to environmental conditions. At the time of +metamorphosis, young are dark brown with specks of black and with a +dark, cuneate, leaflike middorsal mark. The narrow end of this mark +arises just behind the head, and the mark extends posteriorly as far as +the hind leg insertions. At its widest, the mark covers about half the +width of the dorsal surface. The lateral edges of the mark are sharply +defined, but at its anterior and posterior ends it blends into the +ground color. In most individuals smaller than 20 mm., this dorsal mark +is well defined and conspicuous. As growth proceeds, however, it becomes +faint. In frogs 19 to 25 mm. long the marks have disappeared. In +individuals of this size the brown ground color is markedly paler than +in those newly metamorphosed, but is darker than in adults. + +In large adults the dorsal coloration is a uniform pale tan, paler on +the average in females than in males. Temperature and moisture both +affect the shade of coloration. In frogs that were partly desiccated, +the color was unusually pale, with a distinctly greenish tint, and at +high temperatures coloration tended to be relatively pale. + +Hecht and Matalas (1946) have described and figured color patterns in +various populations of _Gastrophryne_, demonstrating geographic trends +and helping to clarify relationships. Their account indicates that the +dark dorsal mark present in young of _olivacea_ but not present in +adults, is better developed and longer persisting in other forms. +Specimens of _carolinensis_, presumably adult, are figured which have +the dark middorsal area contrasting with paler color of the sides. The +dark area is seen to consist of dots or blotches of black pigment which +may be in contact producing more or less continuous black areas, or may +be separate and distinct producing a spotted pattern. Pigmentation is +usually most intense along the lateral edges of the dorsal leaflike +mark; the central portion may be so much paler that the effect is that +of a pair of dorsolateral stripes. This latter type of pattern is best +developed in the population of Key West, Florida. Hecht and Matalas did +not consider these insular frogs to be taxonomically distinct, because +only 48 percent of specimens from the Florida keys had the "Key West" +pattern, while 29 per cent resembled _olivacea_ and 23 per cent +resembled _carolinensis_. In the southwestern subspecies (or species) +_mazatlanensis_, recorded from several localities in Sonora and from +extreme southern Arizona, the dorsal pigmentation similarly tends to be +concentrated in dorsolateral bands, but is much reduced or almost +absent, and there is corresponding pigmentation dorsally across the +middle of the thigh, across the middle of the shank, and on the foot. +When the leg is folded, these three dark areas are brought in contact +with each other and with the dorsolateral body mark, if it is present, +to form a continuous dark area, in a characteristic "ruptive" pattern. +Hecht and Matalas found similar leg bars, less well developed, in +certain specimens of _olivacea_ including one from Gage County, +Nebraska, at the northern end of the known geographic range. + + +MOVEMENTS + +Freiburg (_op. cit._: 384) concluded that ant-eating frogs seem to have +no individual home ranges, but wander in any direction where suitable +habitat is present. However, from records covering a much longer span of +time, it became increasingly evident that a frog ordinarily tends to +stay within a small area, familiar to it and providing its habitat +requirements. + +Nevertheless, in all but a few instances the marked frogs recaptured +were in new locations a greater or lesser distance from the site of +original capture. The movements made by these frogs were of several +distinct types: + + 1. Routine day to day movements from shelter to shelter within + the area familiar to the animal, the "home range." + + 2. Shifts from one home range to another; such shifts may have + been either long or short, and may have occurred abruptly or + by gradual stages. + + 3. Travel by adults to or from a breeding pond. In most or all + instances these adults were regularly established in permanent + home ranges, and they often moved through areas unsuitable + as habitat to reach the ponds. + + 4. Movements of dispersal in the young, recently metamorphosed + and not yet settled in a regular home range. + +Usually there was uncertainty as to which types of movements had been +made by the recaptured individuals. Some may have made two or three +different types of movements in the interval between captures. + +On many occasions individuals were found beneath the same rock on two +consecutive days, or occasionally on several successive days. Rarely, +such continued occupancy of a niche lasted several weeks. In 1949, a +frog was found under the same rock on June 4, 6, 26, 27, and July 1, 3 +and 11. This was an immature female, presumably metamorphosed late in +the summer of 1948. During the five weeks period covered by the records, +it grew from 27 mm. to 34 mm. In 1952, another individual was found +under its home rock on June 23 and 30, July 2 and 3, and August 14 and +20. In 1952 a juvenile was found under a rock on May 30, June 4, and +June 17. These three individuals were exceptional in their continued +occupancy of the same niches. Among the hundreds of others recorded, +none was found more than twice in any one place. + +Despite the fact that field work was concentrated on small areas which +were worked intensively, only eight per cent of the frogs recorded were +ever recaptured, and most of those were recaptured only once. Only 13 +individuals yielded series of records, well spaced, in two or more +different years. These few individuals recaptured frequently may not be +typical of the entire population. The low incidence of recaptures +indicates that relatively few of the frogs present on an area at any one +time have been taken. Because of their secretive and subterranean habits +most of the frogs are missed by a collector who searches by turning +rocks, or trapping with pitfalls. Therefore, even though a marked frog +may survive and remain within a radius of a few hundred feet of one +point for months or even years, the chances of recapture are poor. + +One female was caught first as a juvenile on June 8, 1950. On April 24, +1951, when first recaptured, she had grown to small adult size, and was +only 18 feet from the original location. On July 30, 1951, however, she +was recaptured 750 feet away. At a fourth capture on May 21, 1952, she +had shifted 70 feet farther in the same direction. At the final capture +on June 24, 1952, she was approximately 140 feet from both the third and +fourth locations. The sequence of these records suggests that the frog +had already settled in a home range at the time of her first capture in +1950, and that approximately a year later she shifted to a second home +range, which was occupied for the following year, at least. + +[Illustration: FIG. 8. Distances between captures in frogs marked, and +recaptured after substantial intervals including one or more +hibernations. Distances are grouped in 25-foot intervals. For longer +distances the trend is toward progressively fewer records, indicating +that typical home ranges are small.] + +In several instances, after recaptures as far as 400 feet from the +original location, frogs were again captured near an original location, +suggesting that for some individuals, at least, home ranges may be as +much as 400 feet in diameter. + +Figure 8 shows that for movements of up to 400 feet, numbers of +individuals gradually decrease with greater distance. For distances of +more than 400 feet there are comparatively few records. Of the 59 +individuals recaptured after one or more hibernations, only nine had +moved more than 400 feet from the original location. Twenty-five were +recaptured at distances of 75 feet or less. The mean distance for +movement for all individuals recaptured was 72 feet. A typical home +range, therefore, seems to average no more than 75 feet in radius. Of +the 59 individuals recaptured after one or more hibernations, 47 were +adults and probably many of these had made round-trip migrations to the +breeding pond. This was not actually demonstrated for any one +individual, but several were captured in each of three or four different +years near the same location. + +[Illustration: FIG. 9. Distances between captures and elapsed time in +months in marked frogs recaptured. Few records are for distances more +than 400 feet. There is but little tendency to longer movements in those +caught after relatively long intervals.] + +The trend of movements differed in the sexes. Males are more vagile. Of +21 adult males recaptured, none was less than 40 feet from its original +location, whereas six of the 26 adult females were less than 40 feet +away from the original point of capture. Of seven frogs that had +wandered 700 feet or more, five were males. + + +FOOD HABITS + +According to Smith (1934: 503) stomachs of many specimens, from widely +scattered localities in Kansas, contained only large numbers of small +ants. Tanner (1950: 47) described the situation of a frog found on the +Reservation buried in loose soil beneath a flat rock, beside an ant +burrow, where, presumably, the frog could snap up the passing ants +without shifting its position. Anderson (_op. cit._: 21) examined +alimentary tracts of 203 specimens of _carolinensis_ from Louisiana, +representing a year round sample for several different habitats. He +found a variety of small animals including ants, termites, beetles, +springtails, bugs, ear-wigs, lepidopterans, spiders, mites, centipedes, +and snails. Most of these prey animals were represented by few +individuals, and ants were much more numerous than any of the other +groups. Anderson concluded that ants, termites, and small beetles were +the principal foods. He noted that some of the beetles were of groups +commonly found in ant colonies. Tanner reported that in a large number +of the frogs which he collected in Douglas, Riley, Pottawatomie, and +Geary counties, Kansas, the digestive tracts and feces contained only +ants. Wood (1948: 226) reported an individual of _G. carolinensis_ in +Tennessee found under a flat rock in the center of an ant nest. + +Freiburg (_op. cit._: 383) reported on the stomach contents of 52 +ant-eating frogs collected near the Reservation. Ants constituted nearly +all these stomach contents, though remains of a few small beetles were +found. The ants eaten were of two kinds, _Lasius interjectus_ and +_Crematogaster_ sp. The latter was by far the more numerous. + +Although I made no further study of stomach contents, the myrmecophagous +habits of _Gastrophryne_ have come to my attention frequently in the +course of routine field work. Individuals kept in confinement for a day +or more almost invariably voided feces which consisted mainly or +entirely of ant remains, chiefly the heads, as these are most resistant +to digestion. + +Often upon examining frogs I have found ants (_Crematogaster_ sp.) or +their severed heads, attached with mandibles embedded in the skin. To +have been attacked by ants, the frogs must have been in or beside the +ants' burrow systems. Frequently the frogs that were uncovered beneath +rocks were adjacent to clusters of ants or to their nests or travelways, +in a position strategically located to feed upon them, as described by +Tanner. Often the feces of the frogs were found in pitfalls or under +flat rocks. Although these feces were not analyzed, they seemed to +consist mainly or entirely of ant remains. + +The species of _Crematogaster_, which is the chief food of +_Gastrophryne_ in this region, is largely subterranean in habits, and is +extremely abundant. Any flat rock in damp soil is likely to harbor a +colony beneath it. Colonies are situated also in damp soil away from +rocks, beneath almost any kind of debris, and in hollow weed stalks and +decaying wood. Live-traps for small mammals, having nest boxes attached, +almost always were occupied by colonies of _Crematogaster_, if they were +left in the field in warm, humid weather. Occasionally the ants attacked +and killed small mammals caught in such traps. Among the thousands of +kinds of insects occurring on the Reservation, this ant is one of the +most numerous in individuals, one of the most important on the basis of +biomass and provides an abundant food source for those predators that +are ant eaters. Food supply probably is not a limiting factor to +populations of _Gastrophryne_ on the area. + + +PREDATION + +Young copperheads are known to feed upon ant-eating frogs occasionally +(Anderson, 1942: 216; Freiburg, 1951: 378). Other kinds of snakes +supposedly eat them also. The common water snake (_Natrix sipedon_) and +garter snake (_Thamnophis sirtalis_) probably take heavy toll of the +adults at the time they are concentrated at the breeding pools. Larger +salientians may be among the more important enemies of the breeding +adults, the tadpoles, and the newly metamorphosed young. Bullfrogs +(_Rana catesbeiana_) and leopard frogs (_Rana pipiens_) are normally +abundant at the pond on the Reservation. These large voracious frogs +lining the banks are quick to lunge at any moving object, and must take +heavy toll of the much smaller ant-eating frogs that have to pass +through their ranks to reach the water. The newly metamorphosed young +often are forced to remain at a pond's edge for many days, or even for +weeks, by drought and they must be subject to especially heavy predation +by ranid frogs. Even the smallest newly metamorphosed bullfrogs and +leopard frogs would be large enough to catch and eat them. + +As a result of persistent drought conditions in 1952 and 1953, bullfrogs +were completely eliminated from the pond by early 1954. Re-invasion by a +few individuals occurred in the course of the summer; these probably +made long overland trips from ponds or streams that had persisted +through the drought. Leopard frogs reached the pond in somewhat larger +numbers, but their population in 1954 was only a small percentage of +that present in most other years. Notable success in the ant-eating +frog's reproduction in 1954 may have been due largely to the scarcity of +these large ranids at the breeding ponds. + +Freiburg (_loc. cit._) noted that many of the ant-eating frogs he +examined were scarred, and some had digits or limbs amputated. He did +not speculate concerning the origin of these injuries. However, it seems +likely that many or all of them were inflicted by the short-tailed shrew +(_Blarina brevicauda_). Five-lined skinks living on the same area were +likewise found to be scarred by bites which I identified (Fitch, 1954: +133) as bites of the short-tailed shrew. This shrew is common on the +Reservation, especially in woodland. Many have been trapped in the +pitfalls. On several occasions when a short-tailed shrew was caught in +the same pitfall with ant-eating frogs, it was found to have killed and +eaten them. Like the frogs, the shrews were most often caught in +pitfalls just after heavy rains. Once in 1954 a shrew was found at the +quarry in a pitfall that had been one of those most productive of frogs. +The bottom of the pitfall was strewn with the discarded remains (mostly +feet and skins) of perhaps a dozen ant-eating frogs. All had been eaten +during one night and the following morning, as the trap had been checked +on the preceding day. On other occasions shrews caught in pitfalls with +several frogs had killed and eaten some and left others unharmed. + + +SUMMARY + +In northeastern Kansas the ant-eating frog, _Gastrophryne olivacea_, is +one of the more common species of amphibians. This area is near the +northern limits of the species, genus, and family. The species prefers a +dry, rocky upland habitat often in open woods or at woodland edge where +other kinds of salientians do not ordinarily occur. It is, however, +tolerant of a wide variety of habitat conditions, and may occur in river +flood plains or cultivated land. In these situations where surface rocks +are absent, cracks and rodent burrows presumably furnish the +subterranean shelter that it requires. + +This frog is secretive and spends most of the time in subterranean +shelter, obtaining its food there rather than in the open. Only on warm +rainy nights is it inclined to venture into the open. Then, it moves +about rapidly and with a scuttling gait, a combination of running and +short hops. However, it may be flushed in daylight from a hiding place +by the vibrations from footsteps of a person or an animal, or it may +move about in the daytime when temperatures at night are too low for +activity. Though not swift of foot, the frogs are elusive because of +their tendency to keep under cover, their slippery dermal secretion, and +the ease with which they find and enter holes, or crevices to escape. + +Breeding occurs at any time from late May through August and is +controlled by the distribution of rainfall. Heavy precipitation, +especially rains of two inches or more, stimulates the frogs to migrate +in large numbers to breeding ponds. Even though there are several well +spaced periods of unusually heavy rainfall in the course of a summer, +each one initiates a new cycle of migration, mating and spawning. Heavy +rainfall is a necessity, not only to ensure a water supply in temporary +pools where the frogs breed, but to create the moist conditions they +require for an overland migration. An individual male may migrate to a +pond and breed at least twice in the same season. Whether or not the +females do likewise is unknown. Amplexus and spawning occur mainly +within a day or two after the frogs reach the ponds. The males call +chiefly at night, but there may be daytime choruses when breeding +activity is at its peak. Many males concentrate within a few square +yards in the choruses and float upright usually beside or beneath a stem +or leaf, or other shelter, rendering them extremely inconspicuous. The +call is a bleat of three seconds duration, or a little more. In amplexus +the members of a pair sometimes become glued together by their viscous +dermal secretions. The eggs hatch in approximately 48 hours. The +tadpoles metamorphose in as few as 24 days. Newly metamorphosed frogs +are 15 to 16 mm. in length, or, rarely as small as 14.5 mm. They are +thus much larger than newly metamorphosed _G. carolinensis_, which have +been described as 10-12 mm. or even as small as 8.5 mm. The newly +metamorphosed frogs disperse from the breeding ponds as soon as there is +a heavy rain. The young grow a little more than one mm. in length per +week. Those metamorphosed in early summer may attain minimum adult size +before hibernation which begins in October. It seems that sexual +maturity is most often attained in the second season, at an age of one +to two years. + +_Gastrophryne_ belongs to a family that is primarily tropical in +distribution, and frogs of this genus have much higher temperature +thresholds than most other amphibians of northeastern Kansas, with a +correspondingly short season of activity. For more than half the year, +mid-October to early May the frogs are normally in hibernation. Body +temperatures of active frogs ranged from 17.0 deg. C. to 37.6 deg. C., +but more than two-thirds were within the relatively narrow range, 24.0 +deg. to 31 deg.. Near the date of the first autumn frost the frogs +disappear from the soil surface and from their usual shelters near the +surface, presumably having retired into hibernation in deep holes and +crevices. + +The natural enemies include young of the copperhead. The bullfrog and +leopard frog probably take heavy toll of both the adults and the newly +metamorphosed young at the breeding ponds. Reproductive success of the +ant-eating frogs was much greater in 1954 when these ranids were +unusually scarce. The short-tailed shrew is an important enemy. On +occasion it took heavy toll of frogs trapped in pitfalls, and many of +the larger adults were scarred or mutilated from bites, probably of the +shrew. + +Each of several frogs was found consistently under the same rock for +periods of weeks. The hundreds of other frogs that were marked were +rarely found twice in any one spot. Usually an individual recaptured +after weeks or months was still near the original site. In many +instances the distance involved was only a few yards, but there is some +evidence that home ranges may be as long as 400 feet in greatest +diameter. Of those caught in two or more different years only 15 per +cent were shown to have moved more than 400 feet. These few +exceptionally long movements, up to 2000 feet, involve shifts in home +range or migrations motivated by reproductive urge. + + +LITERATURE CITED + + ANDERSON, P. + 1942. Amphibians and reptiles of Jackson County, Missouri. Bull. + Chicago Acad. Sci., 6: 203-220. + + ANDERSON, P. K. + 1954. Studies in the ecology of the narrow-mouthed toad, Microhyla + carolinensis carolinensis. Tulane Studies in Zool., 2: 15-46. + + BLAIR, A. P. + 1950. Note on Oklahoma microhylid frogs. Copeia, 1950: 152. + + BOGERT, C. M. + 1949. Thermoregulation in reptiles, a factor in evolution. + Evolution, 3: 195-211. + + BRAGG, A. N. + 1943. Observations on the ecology and natural history of Anura, XV. + The hylids and microhylids in Oklahoma. Great Basin Nat., + 4: 62-80. + + de CARVALHO, A. L. + 1954. A preliminary synopsis of the genera of American microhylid + frogs. Occas. Papers Mus. Zool. Univ. Michigan, no. 555: 19 + pp., 1 pl. + + DICE, L. R. + 1923. Notes on the communities of vertebrates of Riley County, + Kansas, with especial reference to the amphibians, reptiles + and mammals. Ecology, 4: 40-53. + + FITCH, H. S. + 1954. Life history and ecology of the five-lined skink, Eumeces + fasciatus. Univ. Kansas Publ. Mus. Nat. Hist., 8: 1-156. + + FREIBURG, R. E. + 1951. An ecological study of the narrow-mouthed toad (Microhyla) in + northeastern Kansas. Trans. Kansas Acad. Sci., 54: 374-386. + + HECHT, M. K., and MATALAS, B. L. + 1946. A review of the Middle American toads of the genus Microhyla. + American Mus. Novitates, no. 1315: 1-21. + + LOOMIS, R. B. + 1945. Microhyla olivacea (Hallowell) in Nebraska. Herpetologica, 2: + 211-212. + + MITTLEMAN, M. B. + 1950. Miscellaneous notes on some amphibians and reptiles from the + southeastern United States. Herpetologica, 6: 20-24. + + PARKER, H. W. + 1934. A monograph of the frogs of the family Microhylidae. British + Mus. (Nat. Hist.) London, vii + 208 pp., figs. 1-67. + + POPE, C. H. + 1931. Notes on amphibians from Fukien, Hainan, and other parts of + China. Bull. American Mus. Nat. Hist., 61: 397-611. + + SCHMIDT, K. P. + 1953. A check list of North American amphibians and reptiles. Univ. + Chicago Press, viii + 280 pp. + + SMITH, H. M. + 1934. The amphibians of Kansas. American Midland Nat., 15: 377-528, + pls. 12-20, maps 1-24. + 1950. Handbook of amphibians and reptiles of Kansas. Univ. Kansas + Publ. Mus. Nat. Hist. Misc. Publ., 2: 1-336 pp., 233 figs. + + SMITH, H. M., and TAYLOR, E. H. + 1950. Type localities of Mexican reptiles and amphibians. Univ. + Kansas Sci. Bull. 33: 313-380. + + STEBBINS, R. C. + 1951. Amphibians of western North America. Univ. California Press, + xviii + 539 pp. + + TANNER, W. W. + 1950. Notes on the habits of Microhyla carolinensis olivacea + (Hallowell). Herpetologica, 6: 47-48. + + WOOD, J. T. + 1948. Microhyla c. carolinensis in an ant nest. Herpetologica, + 4: 226. + + WRIGHT, A. H. + 1932. Life-histories of the frogs of Okefinokee Swamp, Georgia. + Macmillan Co., New York, N. Y. + + WRIGHT, A. H., and WRIGHT, A. A. + 1949. Handbook of frogs and toads of the United States and Canada. + Comstock Publ. Co., Ithaca, New York. + +_Transmitted February 28, 1955._ + + * * * * * + +Transcriber's Notes + + A small number of inconsistencies and typographical errors have been + changed in the text as follows: + + p. 279 "near-by" changed to "nearby" (in nearby counties of Kansas) + p. 289 "successivly" changed to "successively" (two successively older + annual age classes) + p. 297 "per cent" changed to "percent" (only 48 percent of specimens from + the Florida keys) + p. 303 "famliy" changed to "family" (the northern limits of the species, + genus, and family.) + + + + + +End of the Project Gutenberg EBook of A Field Study of the Kansas Ant-Eating +Frog, Gastrophryne olivacea, by Henry S. 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