Bufo terrestris charlesmithi, a New Subspecies from Oklahoma
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1 THE WASMANN JOURNAL OF VOL. 12, No. 2 BIOLOGY S UMJIIER, 1954 Bufo terrestris charlesmithi, a New Subspecies from Oklahoma ARTHUR N. BRAGG,' Unive1 si.ty of Oklahoma, NoTman, Okla.homa About twelve years ago, I discovered a breeding congress of small toads 1.8 miles south and 7 miles east of Norman, Cleveland County, Oklahoma. The animals were breeding in a small cattle tank, dug from the sandy soil in a small clearing on a hillside in the oak-hickory savannah (fig. 1). The breeding call, as first heard from a nearby road, was a long, high-pitched trill which I immediately recorded in notes as that of the American toad, Bufo ten estt is arnet icanus Holbrook; but I also noted that it seemed somewhat higher in pitch than I r emembered for the latter, which at that time I had not heard for several years. At this time, I had not previously seen toads of this type in Oklahoma but had known them all my life in New England. As I emerged f rom the trees near the edge of the pool expecting the American toad, I was quite surprised to find only small individuals. Subsequent search during the next few nights substantiated as fact that the whol e population of these toads in this region were of much smaller individual sizes than most similar individuals of other populations in New England and Wisconsin with which I had formerly been familiar. I have now seen hundreds of these small toads in many parts of eastern Oklahoma and have never yet fo und a really large individual. B ecause of the consistently small size of these toads and the minor difference in their call I suspected that they might represent a new subspecies but I did not think at the time that my experience in the Southwest was sufficient to justify naming as new a population of animals so much like the well-known Ameri- Received for publication May 20, I. Herpetologist. Ok lahoma Biological Survey, and Professor of Zoology, Uni versity of Oklahoma, Norman. [ 245 ]
2 246 THE WASMANN JOURNAL OF BIOLOGY, Vo1.. 12, 1954 can toad. I therefore sought the advice of Mr. Karl P. Schmidt of the Chicago Museum of Natural History and was content, upon securing it, to describe these little toads as a dwarf form of Bufo anwricanus americanus, the name under which the American toad was then generally known (Bragg, 1940, 1940a). Since subsequent observations pr oved the collecting site to be only about 3 to -1 miles east of the western limits of the range of this toad at this latitude in Oklahoma, I interpreted the small size as a case of dwarfing at the western limit of the range. During the years since the first encounter with this form, I have become very familiar with this delicate member of the Bttfo ter- 1 estr is group of toads and have studied and collected examples of it in all parts of eastern Oklahoma. As my experience increased, with it grew the suspicion that my early hunch was right. Finally two influences have convinced me: (1) my study recently of many examples unquestionably belonging to B. t. mnericanus Holbrook from Wisconsin, Indiana, Illinois, New York, New England, and northeastern Kansas in connection with a study of Texas specimens of Bufo (Bragg and Sanders, 1951); (2) H. M. Smith's Figure 1. Type locality for Buto te7testris cha1 lesmithi, 1.8 miles south, 7 miles east of Norman, Cleveland County, Oklahoma, photographed by author on May 9, Eggs were present in the foreground of the pool near the water's edge. Some years age, this pool was deepe ned and its character completely changed.
3 NEW SUBSPECIES OF TOAD FROM OKLAHOMA-BRAGG 247 statement under "Bufo ten est ris" in his handbook (Smith, 1950) which reads: "A distinct subspecies (fig. 48) characterized by small size and unspotted venter occurs in southeastern Kansas but has never been named and properly diagnosed." His figure 48, showing the dorsal side of small toad, might have been made from any one of hundreds of specimens which I have collected, some of them fifteen or less miles from the border of Kansas in the northeastern corner of Oklahoma. I therefore describe this form as new. In naming the new form, I am embarrassed by the necessity of calling it a subspecies, even though I have found no real evidence of intergradation. In itself this would not matter except that I have recently taken a stand that intergradation is "the sine qua non" of a subspecies (Bragg, 1950). I must therefore repudiate this statement (or name the new form as a new species, which it obviously is not). I, therefore, call it a subspecies and propose Bufo terrestris charlesrnithi as its name, in honor of my former colleague, Dr. Charles Chinton Smith, of Pawhuska, Oklahoma. I further suggest "dwarf American toad" as its vernacular name. DIAGNOSIS: A small, Bufo arnericanus-type toad with rough warty back and faintly, if at all, spotted abdomen. It differs from its near relative, B. t. a.rne1 icanus, in consistently smaller size and general lack of mottling on the venter, although some spotting often occurs (see figures 2 and 3). It is also less spiny and usually with no large warts on the hindlegs. DESCRIPTION: Holotype, male, ANB no. 32, collected May 9, 1939 in copulo with female paratype, ANB No. 33. Type locality, 1.8 miles south, 7 miles east of Norman, Cleveland County, Oklahoma. Other paratypes are ANB nos (nos are now Chicago Nat. History Mus. nos ). A description of the holotype as seen alive was published earlier (Bragg, 1940) and is Jwre quoted : "Sex, male; length, tip of the snout to the anterior border of the vent, 59.0 mm.; color, dorsally a yellowish brown, shading to a reddish brown anteriorly. A series of reddish-brown warts, arranged not quite in pairs, each surrounding and often partly covering black spots, which are in turn surrounded by cinnamoncolored areas. Numerous other smaller warts, irregularly scattered, mostly reddish-brown in color, and without black rims. An
4 248 THE WASMANN JOURNAL OF BIOLOGY, VoL. 12, 1954 occasional small black spot or dot, rimmed with cinnamon, elsewhere on the back. A mid-dorsal stripe, whitish in color, extends from about midway between the parotoids to two-thirds the way down the back, widest anteriorly, becoming narrower and less Figure 2. Five representatives of B. t. charlesmithi (about half natural size) to show variation in dorsal characters. Photo by Dr_ Charles C. Smith.
5 NEW SUBSPECIES OF TOAD FROM OKLAHOMA-BRAGG 249 conspicuous posteriorly and with faint indications of it all the way down. Dorsal surface of the thighs with vermiculations of black and with numerous spinous warts of small size, shanks with larger spinous warts, each with a single spine. Cranial crests Figure 3. Buto t. chm lesmithi (slightly enlarged) to show extremes of ventral spotting. The majority of specimens are much less spotted than this. Photo by Dr. Charles C. Smith.
6 250 THE WASMANN JOURNAL OF BIOLOGY, VoL. 12, 1954 conspicuous from the anterior level of the eyes, slightly divergent to the postorbital crests, a spur running mesiad from this point. Post orbital crest and cranial crest make a right angle posterior to the eye on the left, not quite so on the right. Postorbital crests dipping ventrolateral between the eyes and the anterior borders of the tympana, each sending a spur backward to the outer anterior border of the parotoid. Parotoids long, just less than onethird of the length of the body proper; somewhat kidney-shaped and nearest together in the middle. Eyes protuberant, with small warts on the upper eyelid. Tympanum smaller by about one millimeter than the eye, bordered posteriorly by a slight fold of skin. Angle of the mouth slightly posterior to the posterior edge of the tympanum (by about one millimeter), a group of spinous warts between the tympanum and foreleg. Nostrils close tl)gether. Lateral surface colored like the dorsum above, but with black vermiculations in its central and ventral portions, the black extending irregularly upward at the groin and continuing along the posterior surface of the thigh to the vent. Two small reddish warts just cranial of the cloacal opening on the dorsum. Ventral surface granular; background generally light (just off white), but with rather large black irregularly shaped spots from the posterior of the insertions of the arms to half way down the venter. A large elongate black breast-spot, extending from just below the vocal pouch to the posterior level of the insertion of the arms. Throat pouch dusky (not truly black); anterior to this the throat has a 'watery-orange' color which extends also downward along the ventral surface of the lower jaw. Some of this same color on the ventrum of the thighs and on the posterior portion of the belly. Upper jaw with a distinct notch in front, the lower with a corresponding protuberance. Iris distinctly golden but with fine black vermiculations. Two metatarsal tubercles; the outer rounded, the inner larger and with the edge not heavily cornified at the time of description." I have little to add to this except to note the following on ventral coloration, on calling, and on eggs of the new form: In many specimens of B. t. chm lesmithi the venter is spotted, but in others (as noted by H. M. Smith, 1950) it is immaculate. In many having spotted venters, the coloring is quite faint and minor and in none have I seen the heavy mottling characteristic of many s pecimens from the North and East. It would seem, therefore,.
7 NEW SUBSPECIES OF TOAD FROM OKLAHOMA-BRAGG 251 that a tending toward an unspotted venter (complete in some but not in others) is a characteristic of this toad. This, added to its small size, less spininess on the back and legs, and a difference in voice noted below will distinguish this subspecies from Bufo terrestris Bon. and B. t. americanus Holbrook at least as clearly as B. woodhousi woodhousi Girard may be told from B. w. fowleri Hinckley (see H. M. Smith, 1934); and, perhaps more clearly. To determine whether the breeding call of the new form is actually different from that of B. t. ame1 icanus, as my earliest notes on it indicate, I transcribed the calls of B. t. terrestris and B. t. americanus from the Cornell records known as Voices of the Night to a sound scriber disc upon which I also transcribed recordings of B. t. charlesrnithi made by me in Oklahoma. By playing this record I could then compare the calls directly. The result was clear-cut and unmistakable. The call of B. t. charlesmithi is about intermediate in pitch to the calls of the other two, i.e., approximately a half octave lower than B. t. terrest1 is and a half octave higher than B. t. americanus (as estimated, not measured ). The egg-strings are puzzling. I have a distinct recollection that many egg jellies of B. t. charlesmithi that I have seen were indistinguishable from those of B. t. ame1 ica.nus and so state in an earlier publication (Bragg, 1940). However, I find no statement to this effect in my extensive notes; and when I examined three different clutches of preserved eggs, their jellies showed a marked difference from the condition in B. t. ame1 icanus. The difference was that I could find no evidence of "partitions" between adjacent eggs caused by crowding of egg capsules. In fact, I could determine no inner capsule whatever, even with staining; yet, wh en a string was broken between two eggs, the bulging jelly at the break suggested a capsule about each egg inside a continuous tube. These were eggs (in two cases young embryos) preserved in formalin and they appeared perfectly normal. I have had no opportunity to check these observations on live eggs. Further clarification awaits such opportunity. There is little likelihood of my mistaking the species of these particular strings of eggs, for one of them was collected from a pair still producing them in nature and in a r egion of Oklahoma wh ere no other Bufo is known to occur.
8 STATE OF OKLAHOMA T E ~ = ~j:lt llllliiir4tftttllttttl <C Figure 4. Known distribution of B. t. charlesmithi in Oklahoma. Known regions outs ide Okla homa include adjacent.southeastern Arka nsas, Texas just south of J efferson County, Okl ahoma, a nd Ka nsas north of Otta wa County, O){lahoma. 0 ~ "" >-3 II: tg :;J rn > a: > z '-< 0 Cl ~ z > t"' 0 '%j t:li... 0 t"' 0 c;'}.>< < 0 r... ""... <:0 "" ~
9 NEW SUBSPECIES OF TOAD FROM OKLAHOMA- BRAGG 253 I have seen thousands of tadpoles of the new subspecies. In all r espects which I can determine, they are indistinguishable from those of B. t. arne ricanus. The range of B. t. chwrlesrnithi (fi g. 4) probably includes all counties of eastern Oaklahoma and follows the Red River westward to western Love, and Stevens cow1ties and it crosses the river into Texas n ear Nacona. The subspecies also occurs in south western Arkansas but is replaced north of the Arkansas River in Arkansas b:v B. t. arne ricanus, at least in the region of Siloam Springs where I have studied large breeding congresses of this form. On l~ a few miles west of this, in Delaware County, Oklahoma, all of t he Am erican toads seem to be of the dwarf type. Also to the northward in Oklahoma, near the Arkansas line, B. t. charlesmithi alone is known. It also occurs in southeastern Kansas (Smith, 1950) but in northeastern Kansas toads about Lawrence which I examined seemed to be B. t. american us (which is what t h e ~ were labeled at the University of Kansas Museum). Ecologically, B. t. charlesmithi is a fo rm of the oak-hickory savannah, with small populations in the tall-grass prairies adjoining this savannah. For this reason, one might expect this toad in th e Ozarks of Missouri although I have not seen specimens from there. To the south, it apparently does not reach the Red River in southeastern McCurtain County, Oklahoma (where the oak-hickon does not extend) and is thought not to occur in northeastern Texas (Bragg and Sanders, 1951). In Oklahoma, every toad which has been reported since 1935 under the names B. arnericantts, B. americanus ame ricanus, or B. t. ame ricanus may be taken to be B. t. cha.?'lesmithi. It was also B. t. charlesmithi whose food habits were reported (among t hose of other Oklahoma toads) by C. C. Smith and Bragg (1949 ). If B. t. mne1 icanus Holb. occur at all in Oklahoma it must be in the northeast, per h aps in Ottawa County but more likely in Delaware or Adair Counties. Acknowledgments I have worked under the joint auspices of the Department of Zoology, the University Museum, and the Biological Survey of the University of Oklahoma. The Sound Scriber equipment used in recording voices was secur ed by a grant from the Faculty Re-
10 254 THE WASMANN JOURNAL OF BIOLOGY, VoL. 12, 1954 search Committee of the University. Specimens were loaned by the U.S. National Museum (through Dr. Doris M. Cochran), t he University of Illinois Museum of Natural History (through Dr. Hobart M. Smith), and the University of Kansas Museum of Natural History (through Dr. Edward H. Taylor). Specimens were also donated by Mr. Ottys Sanders of the Southwestern Biological Supply Company, Dallas, Texas. Equipment essential to this study was loaned by Dr. Lawrence Poston, Jr. (Department of Modern Languag es) and Dr. Richard Blanc (formerly of the Department of Zoology), both of the University of Oklahoma. I express my appreciation to all t hese people and agen c i e ~->. LITERATURE CITED BRAGG, ARTH UR N Dwarf american toads from Cleveland County, Okla. Proceed Ings Oklahoma Academy of Science, 20: a. Observations on the natural history and ecology of Anura. III. The ecological distribution of Anura of Cleveland County, Oklahoma, including notes on the habits of several species. Ame1 ican Midland Naturalist, 24: BHAGG, A. N., and 0TTYS SANDERS A new subspecies in the Bufo woodho1tsii group of toa ds. The Wasmann Journa l of Biology, 9 : SllnTH, C. C., and A. N. BRAGG Observations on the ecology and natural history of Anura VII. The food a nd feeding habits of the common species of toads in Oklahoma. E cology, 30 : SJ\UTli, H. M The a mphibians of Kansas. American Midland Naturalist, 15: Handbook of amphibians and reptiles of Kansas. Univ e r s it~ of Kansas Museum of Natural History, Miscellaneous Publications, number 2, :!36 pp.
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