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{'.AXONOMY and DISTRIBUTION.OF the SP.ADE.Foor TOADS of NORTH .AMERICA (S.Alienria& PELOBATIDAE)

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{'.AXONOMY and DISTRIBUTION.OF the SP.ADE.Foor TOADS of NORTH .AMERICA (S.Alienria& PELOBATIDAE) ;{'.AXONOMY AND DISTRIBUTION.OF THE SP.ADE.Foor TOADS OF NORTH .AMERICA (S.ALIENrIA& PELOBATIDAE). by Peter S. ghrapliwy A.B. • Universit;sr11oi" K~as, 1953 I ' SUbmi tted to th!?i Tupart:Jimi't C of Zoology and the Faculty of the - Graduate School of the University of Kansas in partial .t'ulf'illment of the requirements £or the de- gree of Master of Arts. TABLE OF CONTEN'l'S Introduction••••••••••••••••••••••••••••••••••••••••••••••••••••• 1 Objectives••••••••••••••••••••••••• .. ••••••••••••••••••••••••••••• 4 Materials and methods•••••• .. •••••••••••••••••••••"••••••••••••••• 4 AoknO"wa-ledgments •••••• ••. •• •••• • •• • ••••••••• •,. •. •· • ., • ~. •• •. •. • •• • • S Taxonomic history ............... • ..............·• ........... • • • • • • • • .. • 7 Chronological list. of specific names ................... •••••••••••••• 11 Sex-limited characters••••••••••••• .. •••••••••• .. •••••••••••••••••• 1.5 Evaluation 0£ taxon0!1lic characters••• .. ••••••••••••••••••••••••••• 16 Artificial key to the genera or American spadefoot toads••••••••• 18 Artificial key to the adults of Genus Soaphiopua ••••••••••••••••• 18 Artificial key to the adults of Genus-~ •••••••••• .. •••••••h•• 19 Genus ~oapbiopug ••••••••••• .. ••••••••••• .. ••••••••••• .. ••••••••••••• 20 Acoounta 0£ species ... •••••••••••••••••••••••••••••••••••••• 20 Soapb,iopp.s holJN:ooki .................................... 20 _a,oa:pbiopus h:µrte;r:i ••••••••••••••••••••••••••••••••••••• JO aeapbi.opu,§ oou~ •••••••••••••••••••••••••••••••••••••• 35 Genus~ •••••••••••••••••••••••••• .. •••••••o•••••• .. ••• .. ••••• .. ••• 49 Account of species and subspecies•••••••••••••••••••••••••• 50 bombitrons •••••••• .. ••••••................................ 50 hammonqa. hammondi ....... •••••o•••o••o•••••.............. 61- ,auea oamm.ondi multiplicata ............................... 71 intermontana ................................................. 81 Phy'logeey ........................................................................ 89 Status of the name Scaphiopidae •••••••••••••••••• .. •••............... 96 SUilllllary and conclusions........................................................... 98 Tables ....................................................... •••••••••••••• .......... 101 Bibliography•••••••••• .. ••• .. ••••• .. •• .. ••••••••• .. •••••• .. • .. •• .. •••• .. •• 110 l. INTRODUCTION Spade.toot toads have been the object of interest for many years but less .is ;yet known about them. than any other group of sallentians of North America. These terrestrial. toadllke a,.,iphi.biana are charaotel"ized by possessing an elliptical pupil, relati~ smooth and thin .troglike skin1 and a. black horny sharp-edged '*spade" on the inner side of ca.oh hind £oot. The latter struoture, ,paralleled in the Genus E,eloba.tu of Europe and Asia, !unctions as an effective burrowing implement. Although more than one hundred years have passed since the first spadefoot was described ·(18.35) by. Harlan from south Carolina, and twelve other living species named since then1 little is yet known concerning the ma.x,y aspects of their We history: This-pre- vails despite the large amount 0£ literature extant on apadefoots. Bragg (1944a, 194.5b) summarized kn01rrledge o:£ the .group and noted the dearth of information about them available to present day herpetologists. Bragg (1944a) notes that because of certain peculiarities manifested by some or all spadefoots, they have £rom the first stimu~ lated great interest. He cites the follmdng possible reasons for this interests "(l) some species are very secretive and shy and are seldom found in numbers and then only a.t'ter long intervals of time; (2) the developmental rate of their embryos and.larvae exceeds that or any connnonly studied amphibian (Wright and Wright, 193.'.H Orten- burger and Ortenburger11 . 1926; Trowbridge and Trowbridge 11 - 19:37; Bragg, 194la) 1 . (3) their distribution is not what might be expected f'rom their structures £or having a thin, rather sort, froglike sld.n suggesting an aquatic or semiaquati.a habitat• all a.re in reality as pearly terrestrial a$ amphibians usua.J.:cy become, and most 0£ the species are limited. to xerl.a habitats in grasslands and deserts I and ( 4) the detail.s of their taxonomic relationships have, till quite recently• been in confusion due in part to lack of specimens, in part to so tew herpetologists having had an opportunity to study them in their native haunts, and in part to a dearth of Wormation on their tadpoles and lite histories." Since spadetoot toads are so imperfectly kn~ man;y- erroneous beliefs have arisen concerning them. Th$ notion most often repeated is that they spend nearly all. of their lives underground and emerge only a.tter heavy rains in spring and summ.er to breed. They then return to their subterranean haunts to awa1t ..turther heavy rains, Bragg (sm,. cit.·) believei, nthe source £or this idea is in the rarity of the solitary or hermit spadefoot of the East, Scaphiopu.s )lglbrooki holbrgoki (Harlan) ••• " As tha West was opened up to settlement other spadef'oots were described. The idea, concerning the rarity 0£ ~• h• holbroolq was applied to these newq described amphibians. Bragg U?Ja• .2.ll,.) mentions he "heard statements clearly . implying that the plains spadefoott a,. P2ffibti(rons Cope, seldom emerges from the ground, and then only once a year to breed." Brooks (193.0) believed that individuals or this species found buried in hard-packed soil in western Oklahoma could not have emerged £or several years. Deck&?" (19:30) reports the discovery in Oklahoma 0£ spadetoots imprisoned in balls of clay several feet belw the surface under conditions dif'£1cu1t to explain. The latter report oertainly- .turthered this belief. Bragg. adds that 0 1ll8.l'ly have accepted this unoritic~, despite the i'aat that it leaves no obvious manner in which they could secure food." Storer (192.511.5:3) also notes that due to the briet spawning period and the shy and secretive nature ot §.. h~ (but true 0£ all other spade£oots) this. belief. became widespread. "But ! ii the observer does not happen to be on band il tbzo right 1im!. and !n th!l! ;.:ight place the animals will $aeape observation entirel;y• tt Considerable evidence is available in the literature to support Storer•s remarks, and to show the fallacy ot this widespread idea. Spade.toot toads abroad and :feeding·in the daytime have been repor-t.ed by a mnnber of naturalists (Gilmore, 1924; Kellogg, l9:32a; Campbell, 1933! Ball, 19.36; Trowbridge and Trowbridge, 19:371 Bragg, 1941, ·1944J Arnold, 19431 Barbour and Gault, 1952; and Pearson, 1955). The generalization that spadei'oots as a group seldom emerge from underground burrows is, therefore, Without foundation in £aot. Newspaper accounts 0£ "raining frogs" ·have found many interested readers. It is -highly probable that. apadef"oot toads have been the source of these stories more often than other amphibians. Spadefoots have no detinite breeding season (Bragg, 194.Sa) and congresses of these animals oocur at nearly any time of year provicling oerta.in climatic conditions prevail (heavy rain, warm tieathar) .-, Under au.oh conditions their sudd$n 11 almost spontaneous emergence would likely lead uninformed observers to report "raining £roga0 •. Abbott (19041 164) and Bragg (1942aal94) have discussed this phenomenon;, OBJECTIVES The aim. o:t the present stu.d;y is to assemble in.one paper the pertinent taxonomic history and distribtltional information 0£ the spade£oot toads ot the Nfrlf. World; point out and correct certain errors o:r nomenclature papetuated by numerousworksrst clarify ' the systematics ot the group; provid~ reatlj- means £or recognizing named :f'orms and; to map the known geographic d:1.st:ribu.tion of eaob. kind. The principal pa.rt. of this work is· dfreoted toward cl~ the taxonomic status o:t those species within the "rumimsmd;ttt group• and emphasiti.ng the taxonontv 0£ the mtom myj,ti;pUcatus, complex. MATERIALS ANO MfffltODS In the course of this study preserved adult specimens have been a.va.ila.b1e to ma. '.J:n addition numerous skeletons ot each kind have been studied. External measurements were made with a vernie~ calip(llr and al1 were taken to the nearest tenth of a m1l.l.i.Ineter. Measurements taken 9.l.'8 as follows: :total J.emtb• From tip o:£ snout to the most poster1ol" projection of the urostyle• 'the back straightened when naoesaary in order to obtain maximum.l.ength. X@ngth 2!. h!!g.. From tip of snou.t in a straight line to the> posterior edge of' the eym.pa.nio membrana. Initi:nar:i.al. dis:t009e .. JJrom modi.an edge of one ~al 'naris to the other. Interorbital distance. Taken transversely where convex surfaces 0£ orbits approached one another most closely. l@lg;th 91. femur. From posterior edge of urostyl.e to distal end. of femur, the limb held at rlght angles to the boey. Itepgt}l gt tibia• From OQnvex surface 0£ knee to convex surtaee . ... .. of heel, the limb oo:tng :f'lexed • . Wj.dth:-2.t ·m,a.4e. Taken at the base of the_ spade, at point ot attachment. The width· o:f the. haad and length the orbit were also \ . ' . or ! '. • recorded at. the illitiation .of'· the study, but these meatro.rements subsequently proved to be so inacourate inpreaerved material as to be useless• Ari attempt has been made to ·ma1ro· all synonymies complete. In• . some instances, hcn-rever, it was :f'ound impossible to relegate ,· ' . positively eaoh reference to its proper plaoe. This resul.ted from my inability to determina precisely what
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