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+Project Gutenberg's Field Study of Kansas Ant-Eating Frog, 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: Field Study of Kansas Ant-Eating Frog
+
+Author: Henry S. Fitch
+
+Release Date: August 29, 2010 [EBook #33574]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** 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.
+  [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 5½ miles north
+and 2½ 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° C. there is little surface activity but individuals
+that had body temperatures as low as 16° 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° C.
+to 37.6° 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° and 31° C. Probably the preferred temperatures lie within
+this range. The highest body temperature recorded, 37.6° 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° to 30.7° in frogs that were found actually moving
+about. Several with relatively low temperatures, 22° to 17°, 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° 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° 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° 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° 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 ½
+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 Peña 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 14½ 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° C. to 37.6° C., but more
+than two-thirds were within the relatively narrow range, 24.0° to 31°.
+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 Field Study of Kansas Ant-Eating Frog, by 
+Henry S. Fitch
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+
+Project Gutenberg's Field Study of Kansas Ant-Eating Frog, 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: Field Study of Kansas Ant-Eating Frog
+
+Author: Henry S. Fitch
+
+Release Date: August 29, 2010 [EBook #33574]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** 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
+
+
+
+
+
+
+</pre>
+
+
+<div class="trans-note">
+<p class="center"><b>Transcriber's Notes</b></p>
+
+<p>A small number of inconsistencies and typographical errors have been
+changed in the text. These are listed <a href="#Transcribers_Notes">at the end of this book</a>.</p>
+
+<p>The Title page and Verso are in error in stating that the pages run 275 to 306. This should read 276-307.</p>
+
+<p>The caption of <a href="#FIG_5">Figure 5</a> states that the illustration is "a little less than twice natural size".
+This is accurate for the linked image when viewed at approx. 100 dpi with browser display setting 100%.
+The thumbnail image is approximately natural size under the same conditions.</p>
+
+<p><b>Table of Contents:</b></p>
+<ul>
+<li><a href="#INTRODUCTION">Introduction</a></li>
+<li><a href="#HABITAT">Habitat</a></li>
+<li><a href="#BEHAVIOR">Behavior</a></li>
+<li><a href="#TEMPERATURE_RELATIONSHIPS">Temperature Relationships</a></li>
+<li><a href="#BREEDING">Breeding</a></li>
+<li><a href="#DEVELOPMENT_OF_EGGS_AND_LARVAE">Development of Eggs and Larvae</a></li>
+<li><a href="#GROWTH">Growth</a></li>
+<li><a href="#COLOR_AND_PATTERN">Color and Pattern</a></li>
+<li><a href="#MOVEMENTS">Movements</a></li>
+<li><a href="#FOOD_HABITS">Food Habits</a></li>
+<li><a href="#PREDATION">Predation</a></li>
+<li><a href="#SUMMARY">Summary</a></li>
+<li><a href="#LITERATURE_CITED">Literature Cited</a></li>
+</ul>
+</div>
+
+
+
+<p class="center">  <span class="smcap">University of Kansas Publications</span></p>
+
+<p class="center">                     <span class="smcap">Museum of Natural History</span></p>
+
+<p class="center">                Volume 8, No. 4, pp. 275-306, 9 figs. in text</p>
+
+<p class="center">                              February 10, 1956</p>
+
+
+<h1>                                A Field Study<br />
+                        of the Kansas Ant-Eating Frog,<br />
+                            Gastrophryne olivacea</h1>
+
+
+<p class="center">                                     BY</p>
+
+<p class="center">                               HENRY S. FITCH</p>
+
+
+<p class="center">                          <span class="smcap">University of Kansas<br />
+                                  Lawrence</span><br />
+                                    1956</p>
+<hr />
+
+<p class="center">  <span class="smcap">University of Kansas Publications, Museum of Natural History</span></p>
+
+<p class="center">           Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+                              Robert W. Wilson</p>
+
+
+
+<p class="center">                Volume 8, No. 4, pp. 275-306, 9 figs. in text<br />
+                        Published February 10, 1956</p>
+
+
+<p class="center">                       <span class="smcap">University of Kansas</span><br />
+                              Lawrence, Kansas</p>
+
+
+<p class="center">                                 PRINTED BY<br />
+                      FERD VOILAND. JR., STATE PRINTER<br />
+                              TOPEKA, KANSAS<br />
+                                    1956</p>
+
+<p class="center">                                  25-7819</p>
+<hr />
+<p><!-- Page 277 --><span class="pagenum"><a name="Page_277" id="Page_277">[Pg 277]</a></span></p>
+
+
+
+<p class="center" style="font-size:x-large">A FIELD STUDY<br />
+OF THE KANSAS ANT-EATING FROG,<br />
+GASTROPHRYNE OLIVACEA</p>
+
+<p class="center">By</p>
+
+<p class="center">Henry S. Fitch</p>
+
+
+<h2><a name="INTRODUCTION" id="INTRODUCTION"></a>INTRODUCTION</h2>
+
+<p>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.</p>
+
+<p>The Reservation, where my field study was made, is the most
+northeastern section in Douglas County, Kansas, and is approximately
+5&frac12; miles north and 2&frac12; 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. <i>G. olivacea</i>, extending north into
+extreme southern Nebraska (Loomis, 1945: 211), ranges farther
+north than any other American species. In the Old World only
+<i>Kaloula borealis</i> has a comparable northward distribution. Occurring
+in the vicinity of Peiping (Pope, 1931: 587), it reaches approximately
+the same latitude as does <i>Gastrophryne</i> in Nebraska.
+The great majority of microhylid genera and species are confined
+to the tropics.</p>
+
+<p>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<!-- Page 278 --><span class="pagenum"><a name="Page_278" id="Page_278">[Pg 278]</a></span>
+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.</p>
+
+<p>While the present report was in preparation, Anderson (1954)
+published an excellent account of the ecology of the eastern species
+<i>G. carolinensis</i> 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.</p>
+
+<p>The frog with which this report is concerned is the <i>Microhyla
+carolinensis olivacea</i> of the check list (Schmidt, 1953: 77) and recent
+authors. De Carvalho (1954: 12) resurrected the generic name,
+<i>Gastrophryne</i>, for the American species formerly included in <i>Microhyla</i>,
+and presented seemingly valid morphological evidence for
+this plausible generic separation.</p>
+
+<p><i>G. olivacea</i> is obviously closely related to <i>G. carolinensis</i>; 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.</p>
+
+<p>Besides <i>G. olivacea</i> and typical <i>G. carolinensis</i> there are several
+named forms in the genus, including some of doubtful status. The
+name <i>mazatlanensis</i> has been applied to a southwestern population,
+which seems to be a well marked subspecies of <i>olivacea</i>, but as yet
+<i>mazatlanensis</i> has been collected at few localities and the evidence
+of intergradation is meager. The names <i>areolata</i> and <i>texensis</i> have
+been applied to populations in Texas. Hecht and Matalas (1946: 3)<!-- Page 279 --><span class="pagenum"><a name="Page_279" id="Page_279">[Pg 279]</a></span>
+consider <i>areolata</i> to be a synonym of <i>olivacea</i>, applied to a population
+showing intergradation with <i>carolinensis</i>, but Wright and
+Wright (1949: 568) consider <i>areolata</i> to be a distinct subspecies.
+<i>G. texensis</i> generally has been considered to be a synonym of
+<i>olivacea</i>. Other species of the genus include the tropical <i>G. usta</i>,
+<i>G. elegans</i> and <i>G. pictiventris</i>.</p>
+
+<p>Of the vernacular names hitherto applied to <i>G. olivacea</i> 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 <i>Rana</i> and <i>Bufo</i> 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 <i>Engystoma</i>, a synonym
+of <i>Gastrophryne</i>. <i>G. olivacea</i> 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 (<i>mazatlanensis</i>) and a
+more eastern one (<i>carolinensis</i>). The names <i>texensis</i>, <i>areolata</i> and
+<i>carolinensis</i> have all been applied to populations in Texas, and it is
+questionable whether typical <i>olivacea</i> even extends into Texas.</p>
+
+
+<h2><a name="HABITAT" id="HABITAT"></a>HABITAT</h2>
+
+<p>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,<!-- Page 280 --><span class="pagenum"><a name="Page_280" id="Page_280">[Pg 280]</a></span>
+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.</p>
+
+<p>Although <i>G. olivacea</i> 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 <i>Gastrophryne</i> 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 (<i>Geomys bursarius</i>),
+would provide subterranean shelter for the frogs, which are not
+efficient diggers.</p>
+
+<p>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.</p>
+
+<p>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<!-- Page 281 --><span class="pagenum"><a name="Page_281" id="Page_281">[Pg 281]</a></span>
+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 (<i>Microtus ochrogaster</i>)
+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.</p>
+
+<p>Anderson (1954: 17) indicated that <i>G. carolinensis</i> in Louisiana
+likewise occurs in diverse habitats, being sufficiently adaptable to
+satisfy its basic requirements in various ways.</p>
+
+
+<h2><a name="BEHAVIOR" id="BEHAVIOR"></a>BEHAVIOR</h2>
+
+<p>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.</p>
+
+<p>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.<!-- Page 282 --><span class="pagenum"><a name="Page_282" id="Page_282">[Pg 282]</a></span></p>
+
+<p>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° C.
+there is little surface activity but individuals that had body temperatures
+as low as 16° C. have been found moving about.</p>
+
+<p>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.</p>
+
+
+<h2><a name="TEMPERATURE_RELATIONSHIPS" id="TEMPERATURE_RELATIONSHIPS"></a>TEMPERATURE RELATIONSHIPS</h2>
+
+<p>Ant-eating frogs are active over a temperature range of at least
+16° C. to 37.6° 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.</p>
+
+<p>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.</p>
+
+<p>Body temperatures of the frogs were taken with a small mercury
+thermometer of the type described by Bogert (1949: 197); the bulb<!-- Page 283 --><span class="pagenum"><a name="Page_283" id="Page_283">[Pg 283]</a></span>
+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.</p>
+
+<div class="figcenter" style="width: 400px;">
+<a href="images/i_009.jpg"><img src="images/i_009_tn.jpg" width="400" height="346" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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° and 31° C. Probably the preferred
+temperatures lie within this range. The highest body temperature
+recorded, 37.6° 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<!-- Page 284 --><span class="pagenum"><a name="Page_284" id="Page_284">[Pg 284]</a></span>
+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° to
+30.7° in frogs that were found actually moving about. Several with
+relatively low temperatures, 22° to 17°, 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.</p>
+
+<div class="figcenter" style="width: 397px;">
+<a href="images/i_010.jpg"><img src="images/i_010_tn.jpg" width="397" height="400" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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<!-- Page 285 --><span class="pagenum"><a name="Page_285" id="Page_285">[Pg 285]</a></span>
+rain of .55 inches ending many weeks of drought. Air temperature
+had been little above 10° C. that night, but had often been below
+freezing in the preceding five weeks.</p>
+
+<p>Reactions of these same two individuals to low temperatures were
+tested in the laboratory. At a body temperature of 11° 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° 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.</p>
+
+
+<h2><a name="BREEDING" id="BREEDING"></a>BREEDING</h2>
+
+<p>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.</p>
+
+<p>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° 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 &frac12; 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 <i>Polygonum</i> and other weeds.
+There was sporadic calling even in daylight and there was a great<!-- Page 286 --><span class="pagenum"><a name="Page_286" id="Page_286">[Pg 286]</a></span>
+chorus each evening for the next few days, but its volume rapidly
+diminished.</p>
+
+<p>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.</p>
+
+<p class="center">
+<span class="smcap">Table 1. Numbers of Frogs Caught Within Two Days After Rain in Funnel<br />
+Traps in 1954, from Mid-June, to the Time of First Frost.</span></p>
+
+
+
+<div class="center">
+<table border="0" cellpadding="4" cellspacing="0" summary="Numbers of frogs caught. See caption above.">
+<tr><th align="center">Date</th><th align="center">Precipitation <br />in inches</th><th align="center">No. of <br />caught frogs</th></tr>
+<tr><td align="left">July 1</td><td align="right" class="padright">2.02</td><td align="right" class="padright">8</td></tr>
+<tr><td align="left">July 10</td><td align="right" class="padright">.11</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">July 16</td><td align="right" class="padright">1.26</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">July 20-21</td><td align="right" class="padright">.94</td><td align="right" class="padright">3</td></tr>
+<tr><td align="left">July 24</td><td align="right" class="padright">.38</td><td align="right" class="padright">2</td></tr>
+<tr><td align="left">July 28</td><td align="right" class="padright">.29</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">August 1-2</td><td align="right" class="padright">3.22</td><td align="right" class="padright">31</td></tr>
+<tr><td align="left">August 6-7-8</td><td align="right" class="padright">2.43</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">August 12</td><td align="right" class="padright">.28</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">August 16</td><td align="right" class="padright">.29</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">August 19-22</td><td align="right" class="padright">.70</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">August 27-28</td><td align="right" class="padright">1.05</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">September 9</td><td align="right" class="padright">.50</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">September 29-30</td><td align="right" class="padright">.38</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">October 4</td><td align="right" class="padright">.74</td><td align="right" class="padright">none</td></tr>
+<tr><td align="left">October 12-14</td><td align="right" class="padright">3.51</td><td align="right" class="padright">none</td></tr>
+</table></div>
+
+
+<p>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.</p>
+
+<p>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<!-- Page 287 --><span class="pagenum"><a name="Page_287" id="Page_287">[Pg 287]</a></span>
+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.</p>
+
+<p>Having heard the call of typical <i>G. carolinensis</i> in Louisiana, I
+have the impression that it is a little shorter, more sheeplike, and less
+insectlike than that of <i>G. olivacea</i>. The call of <i>Gastrophryne</i> 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&#x0113;&#x0113; followed by a bleat."</p>
+
+<p>Stebbins observed breeding choruses (<i>mazatlanensis</i>) at Peña
+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 (<i>op. cit.</i>:
+34) found that in small groups of calling <i>G. carolinensis</i> 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.</p>
+
+<p>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<!-- Page 288 --><span class="pagenum"><a name="Page_288" id="Page_288">[Pg 288]</a></span>
+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.</p>
+
+<p>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.</p>
+
+<p>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 <i>Gastrophryne</i>. 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
+(<i>op. cit.</i>) who observed many details of the mating behavior of
+<i>G. carolinensis</i>, both in the laboratory and under natural conditions,
+mentioned no such adhesion between members of a pair.</p>
+
+<p>Anderson (<i>op. cit.</i>: 31) discussed the possibility that reproductive
+isolation might arise in sympatric populations, such as those of <i>G.
+carolinensis</i> in southern Louisiana, through inherent differences in
+time of spawning. However, in <i>G. olivacea</i> 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 <i>G. olivacea</i>
+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<!-- Page 289 --><span class="pagenum"><a name="Page_289" id="Page_289">[Pg 289]</a></span>
+sexually mature in the early part of the breeding season, may comprise
+the bulk of the breeding population in late summer.</p>
+
+
+<h2><a name="DEVELOPMENT_OF_EGGS_AND_LARVAE" id="DEVELOPMENT_OF_EGGS_AND_LARVAE"></a>DEVELOPMENT OF EGGS AND LARVAE</h2>
+
+<p>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&frac12; 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.</p>
+
+<p>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.</p>
+
+
+<h2><a name="GROWTH" id="GROWTH"></a>GROWTH</h2>
+
+<p>Little has been recorded concerning the growth rate of <i>Gastrophryne</i>
+or the time required for it to attain sexual maturity. Wright
+(1932) found that <i>G. carolinensis</i> 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.</p>
+
+<p>Allowing for size differences between the two species, Wright's
+and Anderson's conclusions regarding growth in <i>G. carolinensis</i>, on
+the basis of size groups, are largely substantiated by my own data
+on the growth of marked individuals of <i>G. olivacea</i> living under
+natural conditions in Kansas.</p>
+
+<p>In 1954, an opportunity to investigate the early growth was afforded<!-- Page 290 --><span class="pagenum"><a name="Page_290" id="Page_290">[Pg 290]</a></span>
+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.</p>
+
+<p class="center">
+<a name="TABLE_2" id="TABLE_2"></a>
+<span class="smcap">Table 2. Growth in Frogs Metamorphosed in the Last Week of
+August, 1954.</span></p>
+
+
+<div class="center">
+<table border="0" cellpadding="4" cellspacing="0" summary="Growth in frogs. See caption above.">
+<tr><th align="center">Time of sample</th><th align="center">Number<br />in sample</th><th align="center">Mean size<br />in mm.</th><th align="center">Size range<br />in mm.</th></tr>
+<tr><td align="left">August 27 to 31</td><td align="right" class="padright">27</td><td align="center">15.55 ± .079</td><td align="left">15 to 17</td></tr>
+<tr><td align="left">September 11</td><td align="right" class="padright">114</td><td align="center">17.2&ensp; ± .033</td><td align="left">14 to 20</td></tr>
+<tr><td align="left">September 15 to 22</td><td align="right" class="padright">12</td><td align="center">18.7&ensp; ± .090</td><td align="left">16 to 20</td></tr>
+<tr><td align="left">September 27 to 30</td><td align="right" class="padright">37</td><td align="center">19.3&ensp; ± .055</td><td align="left">17 to 21.5</td></tr>
+<tr><td align="left">October 1 to 7</td><td align="right" class="padright">62</td><td align="center">20.8&ensp; ± .072</td><td align="left">17 to 24</td></tr>
+<tr><td align="left">October 12 to 17</td><td align="right" class="padright">49</td><td align="center">22.3&ensp; ± .092</td><td align="left">18 to 24</td></tr>
+</table></div>
+
+
+<p>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.</p>
+
+<p>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<!-- Page 291 --><span class="pagenum"><a name="Page_291" id="Page_291">[Pg 291]</a></span>
+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.</p>
+
+<div class="figcenter" style="width: 285px;">
+<a name="FIG_3" id="FIG_3"></a>
+<a href="images/i_017.jpg"><img src="images/i_017_tn.jpg" width="285" height="400" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+<hr />
+</div><p><!-- Page 292 --><span class="pagenum"><a name="Page_292" id="Page_292">[Pg 292]</a></span></p>
+
+<div class="figcenter" style="width: 378px;">
+<a href="images/i_018.jpg"><img src="images/i_018_tn.jpg" width="378" height="400" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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 <a href="#TABLE_2">Table 2</a> and <a href="#FIG_3">Fig. 3</a>. However,
+many juveniles that were captured and marked within a few
+weeks of metamorphosis were recaptured as adults. The selected
+individuals in <a href="#TABLE_3">Table 3</a> are considered typical of growth from "half-grown"
+to small adult size. Growth in many other individuals is
+shown in Figs. <a href="#FIG_6">6</a> and <a href="#FIG_7">7</a>.<!-- Page 293 --><span class="pagenum"><a name="Page_293" id="Page_293">[Pg 293]</a></span></p>
+
+<p class="center">
+<a name="TABLE_3" id="TABLE_3"></a>
+<span class="smcap">Table 3. Growth in Frogs Marked as Young and Recaptured as Small<br />
+Adults.</span></p>
+
+
+
+<div class="center">
+<table border="0" cellpadding="4" cellspacing="0" summary="Growth in marked and recaptured frogs. See caption above.">
+<tr>
+<th align="center" class="bt">Individual <br />and sex</th>
+<th align="center" class="bt">Dates <br />of capture</th>
+<th align="center" class="bt">Length <br />in mm.</th>
+<th align="center" class="bt">Probable time <br />of metamorphosis</th>
+</tr>
+<tr>
+<td align="left" class="bt">No. 1 &#x2640;</td>
+<td align="left" class="bt">August 28, 1951</td>
+<td align="left" class="bt">21.5</td>
+<td align="left" class="bt">Mid-July, 1951</td>
+</tr>
+<tr>
+<td align="left"></td>
+<td align="left">May 5, 1952</td>
+<td align="left">23</td>
+</tr>
+<tr>
+<td align="left"></td>
+<td align="left">July 3, 1952</td>
+<td align="left">32</td>
+</tr>
+<tr><td align="left"></td>
+<td align="left">August 31, 1952</td>
+<td align="left">33</td>
+</tr>
+<tr><td align="left" class="bt">No. 2 &#x2640;</td>
+<td align="left" class="bt">June 8, 1950</td>
+<td align="left" class="bt">25</td>
+<td align="left" class="bt">Late July, 1949</td>
+</tr>
+<tr>
+<td align="left"></td>
+<td align="left">May 24, 1951</td>
+<td align="left">31</td>
+</tr>
+<tr><td align="left"></td>
+<td align="left">July 30, 1951</td>
+<td align="left">34</td>
+</tr>
+<tr><td align="left"></td>
+<td align="left">June 24, 1952</td>
+<td align="left">35</td>
+</tr>
+<tr><td align="left" class="bt">No. 3 &#x2642;</td>
+<td align="left" class="bt">August 31, 1951</td>
+<td align="left" class="bt">24</td>
+<td align="left" class="bt">Late June, 1951</td>
+</tr>
+<tr>
+<td align="left" class="bb"></td>
+<td align="left" class="bb">May 23, 1953</td>
+<td align="left" class="bb" colspan="2">32</td>
+</tr>
+</table></div>
+
+<hr />
+<div class="figcenter" style="width: 327px;">
+<a name="FIG_5" id="FIG_5"></a>
+<a href="images/i_019.jpg"><img src="images/i_019_tn.jpg" width="327" height="244" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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<!-- Page 294 --><span class="pagenum"><a name="Page_294" id="Page_294">[Pg 294]</a></span>
+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.</p>
+
+<div class="figcenter" style="width: 400px;">
+<a name="FIG_6" id="FIG_6"></a>
+<a href="images/i_020.jpg"><img src="images/i_020_tn.jpg" width="400" height="320" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>Differences in size between species and geographic variation in
+size in <i>Gastrophryne</i> have been given little attention by herpetologists,
+but if understood, would help to clarify relationships. Hecht<!-- Page 295 --><span class="pagenum"><a name="Page_295" id="Page_295">[Pg 295]</a></span>
+and Matalas stated in their revision (1946: 5) that size is of no importance
+as a taxonomic character, as typical <i>carolinensis</i>, <i>olivacea</i>,
+and <i>mazatlanensis</i> all averaged approximately the same&mdash;26 to 28
+mm.&mdash;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 <i>G. olivacea</i> 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.</p>
+
+<div class="figcenter" style="width: 400px;">
+<a name="FIG_7" id="FIG_7"></a>
+<a href="images/i_021.jpg"><img src="images/i_021_tn.jpg" width="400" height="328" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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 <a href="#FIG_6">Fig. 6</a>, and they
+made less rapid growth.</span>
+</div>
+
+<p>The population which I studied may be considered typical of
+<i>G. olivacea</i>. 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 <i>olivacea</i> are of approximately
+50 percent greater length than <i>G. carolinensis</i> as reported
+by Wright and Wright (1949: 573) and Anderson (1954: 41). Yet
+Blair (1950: 152) observed that in eastern Oklahoma, where the<!-- Page 296 --><span class="pagenum"><a name="Page_296" id="Page_296">[Pg 296]</a></span>
+ranges of <i>olivacea</i> and <i>carolinensis</i> 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 <i>carolinensis</i> to clasp the smaller females of
+<i>olivacea</i>.</p>
+
+<p>That size differs in different populations, and is still poorly understood,
+is illustrated by the following discrepant figures from various
+authors.</p>
+
+<p class="center"><span class="smcap">Table 4. Size Range of Adults in Various Populations of Gastrophryne.</span></p>
+
+
+
+
+<div class="center">
+<table border="0" cellpadding="4" cellspacing="0" summary="Size range of adult frogs. See caption above.">
+<tr><th align="center">Species or <br />subspecies</th><th align="center">Geographic population <br />sampled</th><th align="center">Authority</th><th align="center">Size range of <br />adults in mm.</th></tr>
+<tr><td align="left"><i>olivacea</i></td><td align="left">Douglas Co., Kansas</td><td align="left">present study</td><td align="left" class="padleft">31 &ensp; to 42</td></tr>
+<tr><td align="left"><i>olivacea</i></td><td align="left">entire range</td><td align="left">Wright and Wright (1949)</td><td align="left" class="padleft">19 &ensp; to 38</td></tr>
+<tr><td align="left"><i>carolinensis</i></td><td align="left">entire range</td><td align="left">Wright and Wright (1949)</td><td align="left" class="padleft">20 &ensp; to 36</td></tr>
+<tr><td align="left"><i>carolinensis</i></td><td align="left">southern Louisiana</td><td align="left">Anderson (1954)</td><td align="left" class="padleft">22 &ensp; to 35</td></tr>
+<tr><td align="left"><i>areolata</i></td><td align="left">southeastern Texas</td><td align="left">Wright and Wright (1949)</td><td align="left" class="padleft">23 &ensp; to 29</td></tr>
+<tr><td align="left"><i>mazatlanensis</i></td><td align="left">Arizona and New Mexico</td><td align="left">Wright and Wright (1949)</td><td align="left" class="padleft">22 &ensp; to 30</td></tr>
+<tr><td align="left"><i>mazatlanensis</i></td><td align="left">Santa Cruz Co., Arizona</td><td align="left">Stebbins (1951)</td><td align="left" class="padleft">25.2 to 31.5</td></tr>
+</table></div>
+
+
+
+<h2><a name="COLOR_AND_PATTERN" id="COLOR_AND_PATTERN"></a>COLOR AND PATTERN</h2>
+
+<p>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.</p>
+
+<p>In large adults the dorsal coloration is a uniform pale tan, paler
+on the average in females than in males. Temperature and moisture<!-- Page 297 --><span class="pagenum"><a name="Page_297" id="Page_297">[Pg 297]</a></span>
+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.</p>
+
+<p>Hecht and Matalas (1946) have described and figured color patterns
+in various populations of <i>Gastrophryne</i>, demonstrating geographic
+trends and helping to clarify relationships. Their account
+indicates that the dark dorsal mark present in young of <i>olivacea</i> but
+not present in adults, is better developed and longer persisting in
+other forms. Specimens of <i>carolinensis</i>, 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 <i>olivacea</i> and 23 per cent resembled
+<i>carolinensis</i>. In the southwestern subspecies (or species) <i>mazatlanensis</i>,
+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 <i>olivacea</i> including one
+from Gage County, Nebraska, at the northern end of the known
+geographic range.</p>
+
+
+<h2><a name="MOVEMENTS" id="MOVEMENTS"></a>MOVEMENTS</h2>
+
+<p>Freiburg (<i>op. cit.</i>: 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.<!-- Page 298 --><span class="pagenum"><a name="Page_298" id="Page_298">[Pg 298]</a></span></p>
+
+<p>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:</p>
+
+<ul>
+<li>1. Routine day to day movements from shelter to shelter within
+   the area familiar to the animal, the "home range."</li>
+
+<li>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.</li>
+
+<li>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.</li>
+
+<li>4. Movements of dispersal in the young, recently metamorphosed
+   and not yet settled in a regular home range.</li>
+</ul>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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<!-- Page 299 --><span class="pagenum"><a name="Page_299" id="Page_299">[Pg 299]</a></span>
+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.</p>
+
+<p>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.</p>
+
+<div class="figcenter" style="width: 400px;">
+<a name="FIG_8" id="FIG_8"></a>
+<a href="images/i_025.jpg"><img src="images/i_025_tn.jpg" width="400" height="279" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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.</p>
+
+<p><a href="#FIG_8">Figure 8</a> shows that for movements of up to 400 feet, numbers of
+individuals gradually decrease with greater distance. For distances<!-- Page 300 --><span class="pagenum"><a name="Page_300" id="Page_300">[Pg 300]</a></span>
+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.</p>
+
+<div class="figcenter" style="width: 400px;">
+<a href="images/i_026.jpg"><img src="images/i_026_tn.jpg" width="400" height="267" alt="" title="" /></a>
+<span class="caption"><span class="smcap">Fig.</span> 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.</span>
+</div>
+
+<p>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.</p>
+
+
+<h2><a name="FOOD_HABITS" id="FOOD_HABITS"></a>FOOD HABITS</h2>
+
+<p>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<!-- Page 301 --><span class="pagenum"><a name="Page_301" id="Page_301">[Pg 301]</a></span>
+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 (<i>op. cit.</i>:
+21) examined alimentary tracts of 203 specimens of <i>carolinensis</i> 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 <i>G. carolinensis</i> in Tennessee found
+under a flat rock in the center of an ant nest.</p>
+
+<p>Freiburg (<i>op. cit.</i>: 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, <i>Lasius interjectus</i>
+and <i>Crematogaster</i> sp. The latter was by far the more
+numerous.</p>
+
+<p>Although I made no further study of stomach contents, the myrmecophagous
+habits of <i>Gastrophryne</i> 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.</p>
+
+<p>Often upon examining frogs I have found ants (<i>Crematogaster</i>
+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.</p>
+
+<p>The species of <i>Crematogaster</i>, which is the chief food of <i>Gastrophryne</i>
+in this region, is largely subterranean in habits, and is extremely
+abundant. Any flat rock in damp soil is likely to harbor<!-- Page 302 --><span class="pagenum"><a name="Page_302" id="Page_302">[Pg 302]</a></span>
+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
+<i>Crematogaster</i>, 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 <i>Gastrophryne</i>
+on the area.</p>
+
+
+<h2><a name="PREDATION" id="PREDATION"></a>PREDATION</h2>
+
+<p>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
+(<i>Natrix sipedon</i>) and garter snake (<i>Thamnophis sirtalis</i>) 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 (<i>Rana catesbeiana</i>) and leopard
+frogs (<i>Rana pipiens</i>) 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.</p>
+
+<p>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.</p>
+
+<p>Freiburg (<i>loc. cit.</i>) noted that many of the ant-eating frogs he<!-- Page 303 --><span class="pagenum"><a name="Page_303" id="Page_303">[Pg 303]</a></span>
+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 (<i>Blarina brevicauda</i>). 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.</p>
+
+
+<h2><a name="SUMMARY" id="SUMMARY"></a>SUMMARY</h2>
+
+<p>In northeastern Kansas the ant-eating frog, <i>Gastrophryne olivacea</i>,
+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.</p>
+
+<p>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.</p>
+
+<p>Breeding occurs at any time from late May through August and<!-- Page 304 --><span class="pagenum"><a name="Page_304" id="Page_304">[Pg 304]</a></span>
+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
+<i>G. carolinensis</i>, 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.</p>
+
+<p><i>Gastrophryne</i> 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° C. to
+37.6° C., but more than two-thirds were within the relatively narrow
+range, 24.0° to 31°. 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.<!-- Page 305 --><span class="pagenum"><a name="Page_305" id="Page_305">[Pg 305]</a></span></p>
+
+<p>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.</p>
+
+<p>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.<!-- Page 306 --><span class="pagenum"><a name="Page_306" id="Page_306">[Pg 306]</a></span></p>
+
+
+<h2><a name="LITERATURE_CITED" id="LITERATURE_CITED"></a>LITERATURE CITED</h2>
+
+
+<p class="author">Anderson, P.</p>
+<p class="hangref">    1942. Amphibians and reptiles of Jackson County, Missouri. Bull. Chicago
+          Acad. Sci., 6: 203-220.</p>
+
+<p class="author">Anderson, P. K.</p>
+<p class="hangref">    1954. Studies in the ecology of the narrow-mouthed toad, Microhyla
+          carolinensis carolinensis. Tulane Studies in Zool., 2: 15-46.</p>
+
+<p class="author">Blair, A. P.</p>
+<p class="hangref">    1950. Note on Oklahoma microhylid frogs. Copeia, 1950: 152.</p>
+
+<p class="author">Bogert, C. M.</p>
+<p class="hangref">    1949. Thermoregulation in reptiles, a factor in evolution. Evolution, 3:
+          195-211.</p>
+
+<p class="author">Bragg, A. N.</p>
+<p class="hangref">    1943. Observations on the ecology and natural history of Anura, XV.
+          The hylids and microhylids in Oklahoma. Great Basin Nat., 4: 62-80.</p>
+
+<p class="author">de Carvalho, A. L.</p>
+<p class="hangref">    1954. A preliminary synopsis of the genera of American microhylid frogs.
+          Occas. Papers Mus. Zool. Univ. Michigan, no. 555: 19 pp., 1 pl.</p>
+
+<p class="author">Dice, L. R.</p>
+<p class="hangref">    1923. Notes on the communities of vertebrates of Riley County, Kansas,
+          with especial reference to the amphibians, reptiles and mammals.
+          Ecology, 4: 40-53.</p>
+
+<p class="author">Fitch, H. S.</p>
+<p class="hangref">    1954. Life history and ecology of the five-lined skink, Eumeces fasciatus.
+          Univ. Kansas Publ. Mus. Nat. Hist., 8: 1-156.</p>
+
+<p class="author">Freiburg, R. E.</p>
+<p class="hangref">    1951. An ecological study of the narrow-mouthed toad (Microhyla) in
+          northeastern Kansas. Trans. Kansas Acad. Sci., 54: 374-386.</p>
+
+<p class="author">Hecht, M. K., and Matalas, B. L.</p>
+<p class="hangref">    1946. A review of the Middle American toads of the genus Microhyla.
+          American Mus. Novitates, no. 1315: 1-21.</p>
+
+<p class="author">Loomis, R. B.</p>
+<p class="hangref">    1945. Microhyla olivacea (Hallowell) in Nebraska. Herpetologica, 2:
+          211-212.</p>
+
+<p class="author">Mittleman, M. B.</p>
+<p class="hangref">    1950. Miscellaneous notes on some amphibians and reptiles from the
+          southeastern United States. Herpetologica, 6: 20-24.</p>
+
+<p class="author">Parker, H. W.</p>
+<p class="hangref">    1934. A monograph of the frogs of the family Microhylidae. British Mus.
+          (Nat. Hist.) London, vii + 208 pp., figs. 1-67.</p>
+
+<p class="author">Pope, C. H.</p>
+<p class="hangref">    1931. Notes on amphibians from Fukien, Hainan, and other parts of
+          China. Bull. American Mus. Nat. Hist., 61: 397-611.</p>
+
+<p class="author">Schmidt, K. P.</p>
+<p class="hangref">    1953. A check list of North American amphibians and reptiles. Univ.
+          Chicago Press, viii + 280 pp.</p>
+
+<p><!-- Page 307 --><span class="pagenum"><a name="Page_307" id="Page_307">[Pg 307]</a></span></p>
+
+<p class="author">Smith, H. M.</p>
+<p class="hangref">    1934. The amphibians of Kansas. American Midland Nat., 15: 377-528,
+          pls. 12-20, maps 1-24.</p>
+<p class="hangref">    1950. Handbook of amphibians and reptiles of Kansas. Univ. Kansas
+          Publ. Mus. Nat. Hist. Misc. Publ., 2: 1-336 pp., 233 figs.</p>
+
+<p class="author">Smith, H. M., and Taylor, E. H.</p>
+<p class="hangref">    1950. Type localities of Mexican reptiles and amphibians. Univ. Kansas
+          Sci. Bull. 33: 313-380.</p>
+
+<p class="author">Stebbins, R. C.</p>
+<p class="hangref">    1951. Amphibians of western North America. Univ. California Press, xviii
+          + 539 pp.</p>
+
+<p class="author">Tanner, W. W.</p>
+<p class="hangref">    1950. Notes on the habits of Microhyla carolinensis olivacea (Hallowell).
+          Herpetologica, 6: 47-48.</p>
+
+<p class="author">Wood, J. T.</p>
+<p class="hangref">    1948. Microhyla c. carolinensis in an ant nest. Herpetologica, 4: 226.</p>
+
+<p class="author">Wright, A. H.</p>
+<p class="hangref">    1932. Life-histories of the frogs of Okefinokee Swamp, Georgia. Macmillan
+          Co., New York, N. Y.</p>
+
+<p class="author">Wright, A. H., and Wright, A. A.</p>
+<p class="hangref">    1949. Handbook of frogs and toads of the United States and Canada.
+          Comstock Publ. Co., Ithaca, New York.</p>
+
+
+<p><i>Transmitted February 28, 1955.</i></p>
+
+<div class="trans-note">
+<p class="center"><b><a name="Transcribers_Notes" id="Transcribers_Notes"></a>Transcriber's Notes</b></p>
+
+<p>A small number of inconsistencies and typographical errors have been
+changed in the text:</p>
+
+<ul>
+<li>p. 279 "near-by" changed to "nearby" (in nearby counties of Kansas)</li>
+<li>p. 289 "successivly" changed to "successively" (two successively older annual age classes)</li>
+<li>p. 297 "per cent" changed to "percent" (only 48 percent of specimens from the Florida keys)</li>
+<li>p. 303 "famliy" changed to "family" (the northern limits of the species, genus, and family.)</li>
+</ul>
+</div>
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Field Study of Kansas Ant-Eating Frog, by 
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+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. Fitch
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