{'.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).

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 degree 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 s'.P$()ies was being treated by the author• A ¢omma bas been inserted between the specific name and the author o~ when that persol.l.is not the autho,: of' the name •.

AOKNOW1EDGMENI'S study was oompleted o~ tlU'oughtbe generous loan of specimens, books, and information by a number of individuals. and institutions. I wish to express trl9' indebtedness and sincGre gratitud& to Professot Edw'ard H. Taylor under whose direction this. study has been carried out. His frien~ counsel dur-ir..g the pr9greas . of' this work and generous loan 0£ speo:L'r/lens from his private collection ·and books :from bis library have been o:t invalua.l:>le aide I thank Dr. Doris M. Ooobran o:t the United.States National Musmun 6.

and Dr. Bryce c. Brown or the streoker Museum, Baylor University, £or supplying much need$d information; Professor Norman Hart-weg,

Dr. William E. Duellman, and Mr. George B. Rabb, University of Michigan Museum of Zoology for their hospitality while I t,ra,a there, as well as for tho loan of specimens; Dl"'• Robert F. Inger and Mr. Hymen Marx, Ohioago Natural History Museum, tor numerous courtesies i Professor William B. Da.'Vi.s, .Agricu1tural and Mecha.nical College of Texas, Professor Georges. Myers and Mr. Alan LeViton,, Natural History- Museum, Stanford University, and Professor Robert c. Stebbins,

Museum of Vertebrate Zoology, University of Qal:ii"o:rnia.1 tor the loan of specimens in their care; Professor Hobart M. SD'd,th, University of

Illinois Museum of Natural :History, :tor his stimulating interest• as well as £or the loan of part or the oollection of the MuseumJ Drs.

Charles J. stine, Jr., Baltimore, Maryland and Frederick A. Shannon•

Wickenburg, .Arizona 0 and Professor James s. Findley, Albuquerque, Ner..r Mexico• tor loan of speoimens ;,t;rom their private collections; and to Professors Arthur N•. Bragg and w. Frank Blair :for certain information pertinent to this study. Katherine Buchanan generously presented a large collection of adult and larval spadof'oot toads to ths Um.versity or Kansaa Museum

of MaturaJ. History. The Museum is indeed :tort.unate to have this material in its collections.

To P:rofessor E. Raymond Rall who kindly read the manuscript and

offered helpful criticisms and suggestions, I wish to express my sincere thanks. To m.v w:tf'et Virginia Chrapliwy, I u,,re a debt of gratitude t9.r constant encouragement and cheerful assistance in a multitude oi' ways. Abbrev.tat.ions used in the locality data a:ra as follwss

ON Cbj.oago Natural History Museum

,,( ' CJS P1'1vate collection of Dr., Charles J. stine• Jr. EHT... BMS Private collaotion 0£ Dr. Edward H. Taylor

FAS Private collection of Dr. F. A. Shannon JSF Privato collt1otion of Dr. James s. f1ndle7 STAN Stanford University Natural Jiistor.v Museum

TCWM Texas Cooperative Wildlita Mtiseum MVZ Museum of' Vertebrate Zoology, Universit7 of California UI Uni.varsity of Illinois Museum o:t Natural History

KU University 0£ Kansas Museum o! Natural History UMMZ Univeraitr. or Michigan Museum. ot ·zoology

TAXONOMIC BISTORP ' Ho1brook established the Genus ~caphigpus in 18)6 and designated a,. poli~ as the- type speoies. How'k\ver, this species had been described on& year earlier by Harlan as boJ.brooldi.. Cope (1866 :81) proposed the Genus :tor §.QaphiQPBS 'qomb!frons . Cope, 1863 and designated bombifrol)l as the tY.Pe species of the new genus., In the s~e paper he assigned. ~oa:phiopu§ hammoru.ij. Baird, 1859 •'\J·~ . and S9apW,opus Jl1l!ltipJ.ioatJ.l§ Cope, 186,3 to the Genus~• The. Oemts soapW,opµs was retained for the species .holb:c,gold Harlan, 18:35 and goµehi Baird, 18,54.

*The use of the name Pelobatidae instead of Sea.phiopidae 1s discussed .··. in a separate seotion of thi:3 paper. 8.

In·hie monumental work on the amphibians 0£ North America.

Cope (1889) recognized two species 1n the Genus ,aoaphio,;myi and two speoi.es (one with three eubspeoios) in the Genus~. The most raoent taxonomic study o:r the spadefoot. toads was that of Tanner (19.39). He red:uoed the two genera of Cope to the

subgenera $)apbiopus and~ 0£ the Genus. Sgap}liouu.s. Tanner's .arrangement hs.s.been followed by the majority of herpetologists. Bryce c. Brawn (19.50140). however; recognizes as a Genus distinct £rom §paphio;e:.is. noting that "th& cha:r;-acters .recently pointed out b:v" Tanner and Bragg which sepa.l."ate .:trom scaphiopus seem to be of greater significanoe than the differences between ·, many genera now widell1" recogniz$d.1 e. g., .P§eudaori~ and~- For

that reason it S$811J.S justifiable to recognize~ and Soappiopµa as separate genera."

I concur with the views of Brown and have here recognized and §gaphiopus as distinct genera. The following discusa:i.on

serves as the basis for my decision. Cope {l866s8l) separated his new Genus~ .from §ga.phiopua on the condition of the skin of the head.. Re diagnosed. the f'orme:r

genus as having "derm. distinct trom oranium, which ts 9n1Jr ossified

superiorly·1n two superciliaey" ha?-$. •t .. For ScapW,om he notes, "derm involved in the oephalio ossification, whioh Ls complete." Tanner. (1939:S) writes "while this di.£.f'erenoeil as well as others, exists between the two PJ:;Opos$d genera~ §rulpbiows. and~. there are also

many similarities whiob, in my _opinion~ hold tho species under dis• oussion in one genus. " Tanner (129.. ~.) believes that it is 0£ advantage to divide the Genus acaphiopus into two subgenera, SQ,fil?hiopus. and ~, ~.~4 aa noted above, he does. Several recent authors (Firsohein, 19.50J Brown, 1950; Smith, :+950; Tay+or, 1952; Davis and smith• 1953; Langebartel and Smith, '(·1:· ..-· __ .-:--;; 1954) are also 0£ the opinion that the Genus~ is distinct. From the evidence available to me I have no course but to agree with them. Members of the Genus~ have·a. frontoparietal ("frontal" of Eaton, 1942} fontanelle (pl•. I, figs. 41 5, 6); in Scaphio12us the frontoparietal and nasal bc>nea are broad and a f'ossa is never present (pl. I, i"igs. 1; 2, 3),.. In tey" opinion these differences alone ara of suc:,h magnitude almost as to warrant generic distinct... ion (see comments of Brown.above).

The spade is cuneif'orm in ~. but it is sickle-shaped in

~caphiopus. There is a bony webbing between the posterior border of th~ sacral diapopbyses and shaft of the urostyle in~. In Scapbiopus the bony-webbing between the posterior border o:r the

saoral diapophyses and sha£t ot the urostyle is generally absent.

The pr~hallux is composed of two segments in §qaph,1-opµs; the pre- < hallux is unsegmented in ~- The tadpoles or lim! are larger than those of Scaphiopua and develop into, at most, medium size adults. Those of Sgaphiopus are

smaller but develop into larger adults. (Bragg1 194.5a:58). The eggs of are large and light in oolor, those or Scap);p.opus are small and dark 1n color (Bragg, l945as65). In addition, ecological dif'ferenoea obtain between~ and 10.

~phiopµ§. All members ot a.re found in ~eri.c habitats (as . "/..~-- ')/ defined by Bragg, 5m.• cit.) whereas all members of Soaphio'R\Y!, ( exoept couchi )• have more mesic habitats. The species of {except . oouohi) breed only after violent .. rains the amount is ot less significance .than rate of fall. The speeies of ~oaphi9pw1, breed only after heavy rains; the rate·ot fall is of less signif'i• ca.nee than the amount (Bragg• op. ci~.). w. Frank Blair (19.55) in a tecan-tstudy of mating calls ot certain anurans presents additional evidencGJ wort~ of mention.

Be mentions that the call di:f'f'ers equally as much between and Soaphiopus as do the skeletal and other·morphologioal features.

In h&mnondi. and bombifront the call is distinctly trilled, whereas in Scaph:i.om!! couog.t and Scaph;opu,i hurieri the call is not trilled.

Since the goal oi' evolution is adaptation to environment and behavior patterns ara a product of evolution; they tend.·;to be

*Bragg (l945a:66) remarks that spadefoot toads arose in a xer.l.¢ ofi"lironment presumably in soutlrri'estern North .America, and lived there suf.f'icientl.y long to have had the xerio breeding pattern (as defined by Bragg, ,sm. ~.) firmly fixed 1n their heredity. ill species . of and one speoies ot Sqaphi®M (1• •.!• g9uohi) :.remained 1n the xerio environment; tha other species o:t·soaphiopB4 spread toward the caat and north and in so doing changed their habits slightly to be--

come more mesic. Soaphiows pouohi• by remaining in the original xerio environment retained the primitive xer1o bt-eeding pattern of tha species of~. 11.

distributed, like patterns of mox-phology, among organisms ill' a number 0£ ways.

The l'emarks of Bragg.~. .!Y:• (1950i87) adequately aorwey my views. ttit is true that morphologioal charactere should be con• sidered wherever they are· known to aid in understanding the life of the animal and of its phylogenetic development. If habits ,and habit-patterns are also products o:t evolution,. these·should be u.sed with morphology i,n.olassi:f'ying organisms and studying phylogeny, which is• I believe• the goal 0£ taxonomy ...

CERONOlOOICAL LIST OF SPECIFIC NAMES APPLIED TO

NORTH1 .AMERICAN SPADEFOOT TOADS

Rana holbrookli Harlan, 183.5 ti Medioal and physical. r~searohes, orr original memoirs in medicine, SUX'gfJ:17;. physiology, geology, zoology, and comparative anatomy. Philadelphia, 183.5, p. 10.5. printed by Iqdia n. Bailey. Type looalit;n South Carolina..

aoEmhiOpu.s §Olitarius, Holbrook, 1836~ North American herpetology,

1st ed., vol. 11 1836, P• 85, pl. 12~ Type loealitya (South) carolina., Georgia., and Tennessee" Soaph\2P4! oouoWJ, Baird, 1854. Proo. Acad. Nat .. Sci. Phila• delphia., April, 18.54, P• 62. Type looalityi R(o Nazas, Ooahuil.a, and Matamoros, Tamaulipas, Mext.co, colleoted by Lieut. Darius N. 12.

Couch (U .. s. Nat. Mus. nos. 371'.31 3714, 371.5).•

~caphi.op:g.tt hanynondii Baird91 18,59. Report Pacific R. R. SUi.-wv., vol. 10 91 Will:lamson•a l"Oute, pt. 4 91 :no. 4 91 18.$7 (18.59), P• 12, pl..

28, figs. 2a-2d. Type localityt Fort Reading (near Redding, Shast.3: • County),. California, collected by Dr. J. F. Hammond (U. s. Nat. Mus. no. 369.5).

Scapbiopus !ID¼J.tip}iea:!m! Cope, 186.3. · Proo. Acade Nat •. Sci.

Philadelphia, March, 186:31 P• S2. Type looality; Valley 0£ Mexic011 collected b.r Jno. Potts (U. s. Nat. Mus. no. 3694).

Soappio;eus v¢u12.Cppe1 1863. Proo. Aoad. Nat. Soi. Pbiladelpbiari

March1 186:3, p .. .52. Type locality: Cape st. Lucas, Baja California, collected by John Xantus (U .. s. Nat. Mus. no. ,5893). §caphiopu.s rmotitrep.\g Oope1 1863. Proc. Acad. Nat. Soi. Phila• delpbia• March, 1863, P• .53. TY]>$ looalityt · Matamoros, Tamaulipas,

/' and Rio Nazas, CoahuUa, Mexico, colleoted by Lieut. Darius N. Couch (U. s. Nat. Mus. nos. :3714, 3715) •*

*Baird neglected to cite the catalog numbers ot the ootypes in his description of ,a. oouch11• Cope (186,3) rev.1.tted the Ganus and removed two (u. s. Nat. Mus. nos. 3714• 371.5) of Baird's original cotypes or coucbiland used them as the basis for his nw species x:,eotii"renis. In this same paper Cope cited one speoimen (U, s. Nat. Mu.a. no. 3713) as the basis for the diagnosis of coucbil. According to Kellogg (19J2bt21)·none of the types is extant. aganbiopus bombifron§ -Cope, 1863. Proc. -Aoad. Nat. Sci. Phila- delphia., Maroh, 1863, P• 53• Type localit.ya ·FortUnion,· on YJ!asouri River, lat. 48° N., :trom Mr,vE. J. DGnts (U.- s. Nat. Mus. 110. 3704); on Platte, River• 200 lniles west of Fort Kearney, from w. s. ·wood of Lt. Bryan's Expedition (tJ. s. -Nat. Mwa. no. :;.52o)J Llano· Estaoad.o Texas, Capt. Pope'11 Exped.. Collection (U. s. Nat. Miis. no. ~703)• stAAnilis Cope, ·1e7.;. xu Yarrow, 'surv:. w 100th Mer.1.c1., vol. 5, 1875, ·P• .52.5, pl. 2$, · £1.gs. 6-8• Type·· l.ocalityi Al.to dos ' Utas, Nortmrest New Mood.co, ooll.eated. by E. n. Copa· {u. s. Nat. Mus.

eoap}p.o'OU.qe,lb'gp oarma.n,-1877• Proo. Amer. Assoc. Adv.; Sci.,

Buffalo Meeting, 1877• p. 194 .. -Type locality# Kay West.. Flor1da1 aollectecl by Count Pourta.11s. (MCZ no .. ·1453, 12 ciotypes).

Sgaplp.onµa·-dgges;t.i Brocohi, Bull. Soa .. Philom.., ser. 7, vol. :3, no. 1, 1879, P• 23. Type locality-a Mexico, coll.acted by Alfred Dug{s (Mus. Hist. Nat. Paris, no. 281a, two-ootypes). SoaJ?laopµ.a internontanUJ Cope. 1883,, Proo. Acad. Nat. Soi. Philadelphia, 1883,. P• 1,5'. Type loc;ality; Salt Lake City-. Utah and

Pyramid Lake, Nevada. ('1.'yp8 speoimen(s) preSUl1lably lost)., latioeps Cope. 1893• Am&r. Nat • .- vol. 27, Feb. 1893. P• 1.55,

1 fig. 'fypa looalit;,s "between SeYlllour, in nortmest Texas south of the Red River. and Austin." Colleeted b,r Mt-. William L. Black.•

•nr. Doris M.. Coohran informs me fan lltt.) that th& type specimen is not in the u. s. National Museum.• and that it was not,.. 1,on the list (prepared by the late Dr. E., R.. Dunn) of typa specimens in the museum

0£ tha Philadelphia Acad9tt1Y of Natural Soienoes. ::''S!mPhiopYA hl:\rtm\ Strecker• 1910. Proo. Biol. soc. Washington, vol. 23. July 2)• 1910• P• 116, pl. :3, figs. 3-4• Type· local1ty1 3¼ miles east of Waco, Texas, collected by John Kern Strecker, Jr. (Baylor Univ. Coll. no. 4179). •,aqm;,N,ow plio]?atraghus T~lor, 19,36. Ann. Inst. Biol.. Mexico, vol.

?,, no~ 4• 1936, P• .S1S, pl. 11 fig. 1. Type looa.li.t11 Sherman County,

Kansas 11 •'Edson Beds"• Ogallala .tormati.on, Middle Pliocene, collected by H. T.~ Martin and OJ.au~ Hibbard (r. u. Mus. Vert. Paleo. no. 1430). •sgaphiowt styd~rt Taylor. 19,a. Univ. Kansas Soil Bull., -vol. 2.;, no. 18, June 10 1938, P- 408,(:pls. 42-4,5. Type looal.1ty1 Logan County• Kansas (sea. 7, T. 11s., R. )7'«1;.), a.bout a mile east or "Rhino Hill QUB.rry", near the Wallace County line, "Edson Beds"• Midd1e Pliocene, collected by Frank Studer (K. u. Mus. Vert. Paleo. no. 1478). *§caJ?hiopu.s·ant.J.cvms Tqlor, 1941. Univ. Kansas Publs. state 0eo1.

Surv. Kansas Bull., no • .38, pt. 6• July 7. 19411 P• 184, fig. 2. Type locality& Sherman County, Kansas, "Edson Bads",'Ogallala:f'ormation•

Middle Pliocene• colleoteid by David Dunkle and Edward H. TB3lor (Ke u. Mus. Vert. PQJ.so. no. 1!#69). •~oe..im.\otml sU,vgr§gs Ta;rlort 1942. Unive Kansas Soi• Bull., vol,. 289 noit

10• Nov. 15t 1942, P• 202, pl.. 1.5 1 ti.gs. 2At 2Be Type looalitya Meade County, Kansas, Rexroad f'auna, Upper Pliocene, locality .3, about 16 miles southwest 0£ Meade, Meade County, Kansns• colleoted by Claucia Hibbard and party (K. u. Mus. Vert. Paleo. no. 6368). •Neosqgph1ow1 mblrei T&ylor, 1942. Univ. Kansas Soi. Bull., vol., 28, no. 10, Nov. 15, 1942; P• 204; pl:. 1S 0 figa. SA, SB .. Type locality: Meado ' . ' County, Kansas, Rexroad fauna, Upper Pliocene, locality:,, about 16 miles southwest 0£ Meade• Meade Oounty1 Kansas, oolleoted by Claude lii.bbard and party (X. u. Mus. vart. Paleo. no. 6367). * mopelobatea grandtEt Zweifel• 19,56. .Ame:r. Mus. Novitates, no.

1762, Apr. 6, ;t.9.5·5• P• .'.3t .figs. 1 ...7. Type 1ocalit1_1 Pennington County, South Dakota, ear:cy- Ollgoeene; middle part ot ~em 1U8lliber of Chadron f'ormatlcn, ~5 feet above base. m:vid$ between West Fork and Main Fork ot Indian Creek, one-ha.J.f mile sou~h 0£ f?rksJ southeast corner 0£ the SE ¼<>f sect. 34, T. 3 s., R. 12 E. Oolleote

SEX.... t!MITED ORARACTERS

§exual gilnorebi@z Sec:ondafy aex Ol'i..aracters are manitest in spadefoot toads in several ways. Males have a. vooal pouoh• thioker and shorter> fingers, and more extensive trebbing on the bind toot. The latter oharaeter was first: reported (in Sge:oW,P'P9:§ couohi) by Bogert and oliver U94.5s339) • Davis and Smith (19.5:3t14.5) notijd this exten... sive webbing in b:reeding males ot hmong;b ;ma1t,.p11oat9;. In the present stud;r this oha.raoter wa.1 found to~ quite ¢orad.stent in males ot all species of spa.detoots. The males also have gone:rally a darker throat and ohin than females. In addition, males have the first two fingers, and oocam.ona.ll.y the third, covered w.1.tb black nuptial

•~Fossil. 16.

exoresences. The cot'nified. tip of the :ti:f'th toe is not a se»-limited character,

appearing in males and females. :Ct must be mentioned that the inter- orb1tal. bor.gr boss of Spea bombi.frol'l$ is not l.imited to one sex; both ------. ' ' . sexes possess this ob.araater! I found that the sems of~ M,--'lfflOP~ mgl.ttpl;t®,t{a differ in size,, m:a.les averaging "8.8 •• in anout to vent length, the 1'emal.$s averaging 46.9 mm., in snout.-vent length. The degree ot do:rsa1 t-ugosity in h• mµ.l:ti@oa;w is not a o~aoter of sexual dinlorphism, as suggested by certain individuals. This.is fur&Ml" disoussed under that $'108P8ci•s•

EVALUATION OF .TAXONOMIC O!WtA.CTER.S

The oriteria for speoiation in the North .American P0lobat1ds are based~ upon mo:rphological dif'terences and seoondar:1.l,y upon the color pattern. It seems wise, there.tore, to definia and evaluate the

characters which are most oommonly used in th$ deseriptions and diagno- ses ot spadetoot toads. The di£fere:ntiat.1ng oharaoters ma;v- be divided into £our genera1 groupings o:t dorsum o.f bod;y-, venter of 'body, speoia.lexternal f'eatures. and color pattern. ~ors:um 91. m.

'' ' ha.vi,ng a. rather smooth skin With the tubercles tew and small. faroto~d gJi@:PB:t This gland 1s 1imited to certa:d.n $p$oiea of each

'' genus. Th$ size and shape of the parotoid is ot taxonomic worth in d1s- t1n.guish1ng certain species ~d subspecies of the.Genus.Sq{mbiopus. yentar st~.

' ' fec\QraJ: glandju There are two di.stin<)t glands on th(j ohest of' two SJ>$oies of SCc?zphiopqs. . The £'unction o£ these glands is not Imown.

The small soa.ttered ventral tubercles; usually more concentrated in the inter£emoral. area, ars too variable and are of no value as taxonomic characters. Spgoial: J9.otarnal ieharac~. 'l'ympanuma Present and distinct in onl.y certain species o:r· Sqaphio;g;q,~. It is general.4' indistinct. in all species of ~• Supr~ ~• Because of the variable nature ot these glands in respect to size and number, ·they do not serve as characters of taxonomic worth. Like the pectoral glands the function of these is also not known.

aJaad~s The apadelike process is found in all spade.toot toads.

The shape and size ot the spade in the two genera iB consistent and is of value also in di£.ferentiating the subspecies of~ hanunondi. lllWrorbi tal ~• Present in two species of -™. and two species of Soaph;\opup. The nature of th$ boss, whether bony or glandular• is a distinguishing character in~ bomWJ'rpns and .fL. ~ntermon;tmia. The 18.

degree.of development of the boss.is of taxonomic value in the

distinction of scaphiow, burterj, .· and §.S.aphiopy noibroold.. Color patte;:n. 2olor patterns The color pattern of the head in the diff'erant species ot spadetoot toads shows little indiVidual variation and ;, proves to be a valid criterion. in· this study. Dorsal~ Ratterru Also .aorunatent withl.n ditterent species, it has proved useful in ditterentiating the species namm.ondi and in,termontanq 0£ the Genus ™•

ARTIFIOIAt KEY. TO THE GENERA OF AMERICAN SPADEFOOT TOADS Skin of head coossified with oraniumJ spade sickle-shaped; tympanum and parotoids usually distinct•••••••••••••••••••••• saaehiop;us

Skin o:t head not ooossified with cranium; spade usually rounded1 not sick:ltt-shaped; tympanum and parotoids usually lbldisti:nc~.... ' . . :- ·;

ARTIFICIAL KEY TO THE ADULTS;OFTHE GENUS SCAPHIOPUS

1. Pectoral glands present; body color pattern of two light lines • • 2 ~ectoral glands absent; color pattern appears as a network or marbling, no light lines on dorsum ...... ,?ouob:i 2. Posterior part of interorbital area much elevated and rugooe;

parotoid. usually oval••••••••••••••••••••••••••••••••• hurt§ri Posterior p~ of interorbital area not much elevatedi rarely

rugose; parotoid usuall;r round•••••••••••••••••••••• holbt9ok1 ARTIFICIAL. KEY TO ADULTS OF THE GENUS mA

1. A promin~nt ;S.nterorbital bocy boss (1~ nature evident upon dis• seotion)present ...... bombifrons Interorbital:.bon;y boss absent·•• ...... •• ...... ••• .. ••• ...... •• .... •·•2 2. Do:rsal color pattern of -bro broad light stripes and one light dorsolateral. s,t:r:i.pe; a dark blotch over each upper eyelid passing ·

posteriorly and meeting to::·fornt a V; 1nterorbita1 spa0e often . . ·. . .· . rused pt'Ominent:i.;r._ but :1.s gl.andula:rriand not 'bon;r; spa.de widi.:t%" ··

than long; body usua.l.q i-ugoae .... j ...... i4tgontana ·· . . ' . . . . . Dorsal color pattern not as above. :lnterorbital space not prominentl;r raised, boey relatively smooth or rugosa, parotoid distinct or

J.. Body covered w1. th ;c nume:rous large tuberoles; ·parotoid· present,

ottan enoroacbing upon and ponaealirig tympanUin; ranger south

Body rela.tive'.cy smoothJ parotoidsmall and indistinct, or absentJ tympanum not eoncealedJ ranges California, southtq-esterly to 20.

GENUS SC.APHIOPUS Soaphiopus Holbrook, North .Am.erican Herpetology, 1st ed., vol. 1, 1836, P• 85 (Generic type, soaphiopu¢. so?J;tatj,us Holbrook~. 1836

bolbroold Harlan, 1~ , by mono:typy). Di&aonorua The frontoparietal bones broad, without fontanalle1 skin of head generally ooossifiecl ~th c~anium; skin 0£ dorsum with small. uniform tubercles; spadelike process. sickle-shaped; eggs small and .dark· in . color; tadpoles , small,, usually 2.5 mm. or less in total. length; adults · large, up to 82 mm. in tota1 length; bony webbing be~een posterior border of sacral diapophyses and shaft ot urostyie gener~ absent1 prehall~ segmented ( except in _a. oouchi).

ACCOUNTS OF SPECIES

Seapltj.oJ2U!t holprook:\, (Harlan) (pl. I, fig. 3; pl. n, fig. 4; fig. 1) Holbrooki Harlan, Medical and pbysioal researches, ors original

memoirs in medicine, surgery, physiology, geology, zoology, and

comparative anatomy, 183.5, p. 105. Type locality& South Carolina. soa12hiopu,p solitarius Holbrook, North American Herpetology, 1st ed., vol. 1, 1836, P• 85, pl. 12; Tsohudi, Claesif'. Batrach., 1838, p. 8JJ Dume'ril and Bibron, irpetologie o6'~rale, vol. 8, 1841, p. 473; DeKay-, Zoology or NewYork, vol. 3, 1842, p. 66, pl. 19, fig. 47 (New York); Holbrook, North American Herpetology, 2nd ed., vol. 4,

1642, P• 109, pl. 271 Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 7t 18.55, P• 429 ('ioommon in Georgian);. Gunther, Catalogue of the Batrachia. Salientia in the British Museum. 18.SS, P• 38J Abbott, Ooo1., New Jers~Y,. APl>&Ad:\.Xt, 1869,, p., 804J Garmant Proo~ Amel"~ Assoo. Adv. sci., vol. 2.5, 1877• P• 194 (South caroUna.)1 Bro~cbi~ Mission Soienti:f'ique au Mext.que et .cl.ans l•Amerique Central.et Etudes sur les Batrac:i..ena, livr. 1, 1881• p. 23, pl. 9, figs • .2.. 3 (rtCarolina, Georgia., Mex.t.oo". the latter locality 1-n· errorJ B!o!: mlbrookli placed in ayi'.lo~ of~• eolJ.t&rj.u@}; Boulengerl) Catalogue of the. Batraabia $B.lientia in the British Museum, 2nd ed., 1882, P• 434 (~ holbtpokii and·. §caphiopu,a hql.brooki, placed. into s,norzymy ot

§.. eolit§!tiS§l; Garman, Bull. Essex Inst., vol. 16, 1884, P• 45

(Maasaohu.setts, Florida, Mississippi)J Abbott11 A Naturalists

Rambles About Homs, 1885, p. 476 (habits in New Jersey)J Bou1enger,

Proc. Zool. Soc. London, 1899, p. 790, pl• .52 (redest:ribed and figured); Procter, Proo. Zool. Soc. London, 1921, P• 20.) (disou.ssea

the scapula); Gadow, Amphibia and Reptiles, 1923, P• 164 (general. disoussion) ~. ~h.\omw holbroolg.. Baird, Report Pacilio R. R. Surv•• 10, pt. 4-, no. 4,.

l8.59, P• 12, pl. 28, fig. 1 (figured to compare w:l.th §.• ha.llltllOngi);

Cope, Proc. Acad. Nat. so:1.. Pbiladelpbia,· 18631 p·• .54 (range);

Cope, Jour. Acad. Nat. soi. Phil.a.d&lph1a., vol. 611 ser. 2, 1866,

P• 82 (listed); Putnam, .Amert, Nat., vol .. 11 1867, P• 108 (breeding

behavior in New England)$ Oope 1 u. s .. Nat. Mus. Bull., no. 1, 1875, p. ;32 (range); Coues and Yarro.r. Proc. Acad. Nat. sc1. Philadelphia, 1878, P• 28 (North Carolina), Sid.th,. .Amer. Nat., vol. 13, 1879, P• 651 (Connecticut); Rice and Davi&, Bull. .. Illinois State tab. Nat. Hist., no. 5, 1883, p. 21 (range, "eaatern·u. S.")l Yarrow, Bull. u. s. Nat. Mus •• no. 24, 1883; P• 177 (distribution) J Abbott• Amer. Nat., vol. 18• 1884, p. 1075 (babitt, New Jersey)J Pike• . ': ' ., Bull • .Aln. Mus. Nat. Hist., vol~ 7, 1886, P• 213 _(bx-Ge~ habi.tssi Long Island, New York); Hargitt, .Amer .. Nat., .vol •. 22. 1888, P• 5:35

(Martha's Vin&ya.rd, M.assachuaetts; l>reeding behavio:t")t Cope., Bull.

U. s. Nat. Mus.• no. 34, 1889, P• 298, .i'ig. 74; pl. 57, fig. 2J .· I . pl. 68, fig. 31 pl. 7:;. fig. :30; ~ey, Amax- •. Nat., vol. 30, 1896,. P• 501 (North Carol.ina}i Cope, Amer. Nat., vol. :;o_. 1896·, P• 1011 (geogr. dis:t;.ribution)J Sherwood •. Proo. ~an $oae. New York, 1898~ p. 17 (eggs described); Eckel and Paulmier:, Bull. Netr l'ork State Mus., no. 51, 1902, P• 404 (New York)J Ha;r, Proo. Biol. Soc. Washington, vol. 1.5• 1902, P• 128 (Distx-1¢ of Col\t!llbia); Abbott, Proc. Amel". Philosoph·. Soo., vol. 43. t904, P• 16; (habits h Henshaw, Ocoas. Papers Boston Soc. Nat. Hist • ., vol •. 7,, 1904, P• 8 (looalities in Connecticut and MassachU$ettsh Diokereon, The Frog Book, 1906, P• 53, pls. 6, 7, col. pl. 2 (g~ral. di~cu,ssion)J Fowle?'• Ann. Rept. New Jersey State Mus., 1906, P• 92, pl., 19

(New Jersey)J Hurter and Strecker; Trans. Aead. S?i,• St. 1.,ouis 1 vol. 18, 1909 11 p. 20 (Arkansas); Brim1e;v, Proc. Biol. soo. Washington, ' vol. 23, 1910 1 p. 10 (Florida); Hurter, Trans. Aoad. sci.. St., Louis,

191-1, P• 11L (Missouri hypothetical. list); $tll'i"a.Qa 11 Pennsylvania

Dept. Agrlo. Div. Zool., Bimonthly Zool. Bull.,,:no .. 3, 19131

P• 115, fig. 16 (Pennsylvania); Overton, Bull. Mus. Br~lc:cyn Inst. Arts Sci., vol. 2, no. :3, 1914, p. 28, pls. 5, 6, fig. 39 (Long

Ialand, New' Yorkh Overton, Copeia, no. 20 11 191.5 11 p;. 17 (u:,ng

Island, New York); overton. Oopeia, no. 24,. 1915, p • .52•.S.3 (bNeding habits, New York)J Niohols,. Copeia, no. 4,5, 1917.• P• 59 (Long Island New York); Fowler. Copeia, no. 55, 1918, P• !44. (first report from V!rginia.)j Mellen, .New York Zoo1. Soc., Report Direct.

Aqu.ar., 1919, P• 15 ("nlaXilmnn lite, 6 D19nt~s"H wright and

Wright 11 Oopeia, no. 74, 1919, p. 81 (n«.r county record in New York); Deckert• Copaia, no. 92, 1921, P• 22 (Florida); Dackst-t. Copeia, _no. 112, 1922, P• 88 (Florida raoord)J Nobl.e, :aun._ A.me:r•. Mus. Nat. Hist., vol. 46, 1922, pl. 11 (thigh musculature); Nieden, Das Tierreich, Lie£. LJ6t Anura I, 1923, P• 451 Wright, Oopsia, no. 11.5, 1923, P• :34 (Georgi.ah Wright and wright, Amer. Na.top vol. SS, 1924, P• 379 1 381 1 pl. I (eggs figured.); Br.tmley» Jour. Elisha Mitch&l.1 soi. Soc., vol •. 42• _1926, P• 80 (distribution~ North Oarolinah Brimley, Copeia., no. 162, 1927, P• 11 (additional 2."ecords in North Carolina); Wright.• Protll .. u. s. Nat. Mus.,. vol. 74, 1929, PP• 3, 16• pls. 1, 6 (larvae described and figured); Duntl, Bull. Boston soc. Nat. 1Iist., noo S7, 19:30, P• 6 (Maaaachusetts); Noble, Th& Biology- ot the Amphibia, 19.'.31, pp. 400 (fig. 13.5),

406-407, 409,- 494 (fig. 158) general disouss:ton., ~t and vocal pouah figured); Allen. Oop<,1a1 no .. 2., 19:32; P• 104 .(Mississippi);

Stone; Copeia• l'lOe 11 1932, P• 3_;.,.36 (terrestrial. a.ctiVity, NEn-1 Jersey}; Carr, Florida Nat.,, vol .. ?',~no• 2, 1934, P• 21 (call described)t Driver, Oope1a, no. 1, 1936, PP• 67 ... 69 (breeding habits, early development in Massaohusetts)J Giovannoli, Copaia, no. 1, 19.36,. P• 69 (first report from Kentucky); Ramaswalni• Anat. Anz., Bd., 81, nos. 4-6, 1.936, pp. 6.5 ..96 {crania1 morphology); Br:tmley, Carolina Tips, 1939-194.'.3, unpaged (distribution in North Oarollna); Pawling, He~tologica, vol. 1, no. 6, 19.39, P• 169 (Pennsylvania) s Tanner, Great Basin Nat.,. vol. 1, no. lp 24.

1939, PP• ?-81 pl; 1, figs. 1.;;.24 pl.:, (critical taxonomic study);

Jobson, Herpetologica, ·vol. 2, no. 29 1940; ·p. 40 (Sou.th Carolina);

Green, Pro<'h Wast Virginia Aead. Soi., vol. 201 1948, P• 60 (West Virginia); Smith, Nat. 'Hist. Misc., no. :,j, 1948, P• 2 (seeond report. in Illinois); Engels, Amer .. )11.dl; Nat., vol. 47, no. 3,' 19.52, P• 72.'3 (North Carolilla)J Bbadurit Proc •. Zool. Soc. Bengal. ' vol; 6, 19.53, p~ 20, 'p1.·4, figs. 17.;.21 (µrogem.ta1 sylftem dis--

QUSsed and ttgured)j Endsley, JoUl". Tennessee·Acad.., Soi •• vol. 29,

no~ 1 11 19.54; P• 37 (Tennessae)I Parmalee 11 Ill~nois state Mus.~ 11toi7 of' Illinois ser~, no. 10; 19.54, p. 22 (distribution in Illinois)j Mart.of, Copeia., no.· ;3• 19.5.5& P• 169 (no evidence o£

ooourrenoe in Athens area of Oeorgia)J Auf:f'enbetg, "Breviora, no • .$21 19.56, PP• 6-7 (fossil material from Miocene' of·Florida referred t.o this species). Cs)oaplp.opug §J.p;us Garman, Proc. A:rn&r. Assoc. Adv-. so1., Butf'aJ.o, 25th meeting, 1877, P• 194. Type localltys KeyWest 11 Florida. Sosmhiopus go:J4.tar;tu11 §!:lbu!al• 'Garman, Bull. Essex Inst~, vol. 16~ 1884,.

P• 45 (Key Wast, Flor.1.da). Sgappi9PU§ )lolbtpold. albqs 11 ~jneger and Barbour, Cheak L:utt ot ·North American Amphibians and Reptiles, 1917, ·p. ·26:;. Stejneger and Barbour, p. 23; st~jneger .and Barbour• ~" 3rd ed., ~. 2nd. ed., 1923, .,· 19.3311 P• 26; Wright and Wright, Handbook of Frogs and Toads, 19:33t P• 4:r (general disausaionh Chao Liu~ Proc. Boston soc. Nat. Hi.st .. ; vol. '41, no. 3. 1935, P• :,o (vocal sao'desol'ibed}J au.rt, Trans., Kansas .Adad. sci., vol. 41, i938, p. :335 (Florida); Stejneger and

Barbour, Check List 0£ North .Amerioan J\xilpbibians and Rtptiles1

4th ed., 19)9 1 P• 30; Carr, Univ. Flori4a Publi. Biol.. Soi.,.ser. :3, no. i, 19401 P• 5) (d3.str1.bution in Florida)JWright. .and Wright,

Handbook of Frogs and Toads, 1942, P• $6 (general disouss~on)s

stejneger and Barbour. Check List. 0£ North-.Ame~oa.n. Amphibians and Reptiles, ,5th $d., 19~J, P• :38; Wl"ight and Wright, Handbook of Frogaand Toads ot the United states .and Canada, l.949, PP• 127•. 129, pl. 22 (general discussion); Sc-,bmidt, Check List 0£ North ' American Amphibians @d Reptiles., 6th ed.• 195), P• .58; Duell.man, Oopeia, no. 2, 19.5.5, pp·. 141~14.'.3, fig. 1 (al.J?ps, relegs,ted. to synonymy Qf. ,a. b9Jl?rool4). Soap_}liopu! bolW92lg holb£.9oki, stejneger and Barbour;· Oh$0k List of Iiorth American Alttphibians and Reptiles, 1st ed., 1917, P• 26; Fisher; Copeia• no. 8.5, 1920, P• 76.78 (New Yo:rk)I Stejneger and Barbotll", Check List ot North .American Amphibians and. Reptiles,

2nd ed., 1923, P• 23 (:range)J Va:ri- Hyning, Cor,eia;. no. 118• 1923, P• 68 (Florida); Babcock, Bull. Boston Soc. Nat. Hist., no. :,8,

1926 11 P• 13 (habits in NEY,r England)J Corrington, Oopeia, no. 172, 1929, P• 64~ (South Ca:rol.i.na); Trautman, Copeu, no. 2, 1931,

p. 6.3 (seconctl.reoord. tor Virginia); Stejneger and Barbo\ll", Cheok

Liat of No~h Amer.lean Amphibians and Reptiles, )rd ed., 193J; P• 26 {12m.)1 Van Hyning, Oopaia, nofl lt 193.:3, P• 3 (Florida)I

Wright and Wr:1.ghttBandbook 0£ Frogs and Toads, 19.'.33, P• 42-43 (general discussion); Chao Liu, Proo. Boston Soc. Nat. l!ist.,

vol. 41, no. 3 1 193.5, ·p .. 30 (vooa:L saa desar.1.becih Ta;y1or 11 Univ. Kansas Sci. Bull., vol. 22, no. 10, 193.5, P• 209 (.Arkansas); :sau,- Tl"al>.S• Connecticut Acad. Soi., vol. ;'j2, 1936, PP• :3.51•379, pls. 14, :i.5, figs. 1-2 ( distribution and behavior .in OonneoticutJ r$cords 26.

noted .from Masaaohu.setts and New· York); Brandt, Copeia, no. 4,.

19;6, P• 216. 222 (North Carolina records)J Conant and Bailey,

Oocas. Papers Muth Zool. Univ. Mioh:1.gan, no. 328, 1936, P• 3. (New Jersey); Babbitt, Bull. Connectii:ut· Geol.· Nat. Hist. 'SUl:"V•• no. 57, 1937, P• 26, pl. 10 (distribution in Connecticut); Smith, Herpetologica,vol. 1, 19:37, PP• 104-108 (akul.ltigured.1 nolbrooki distinct from bmi,e~); Black and M1:Jnger, Ocoas. Papers Univ.

• Arkansas Mus., no. 2• 19381 P• 13 (~.) (distribution in Arkansas); Burt, Trans Kansas Aaad. Sci., vol. 41, 1938, P• 335

{records from· Flor:tda.1 Georgia, Mississippi, 1-iorth Clarol.ina, South

Carolina); swanson, Amer. Mi.dl. Nat.; vol. 201 no. 3. 1938, P• 71.'.3

(.first record from Indiana)J •Wright and Wright, Trana. Texas Aoad.

Soi., vol .. 21, ·1938.- P• 18 (hypothetical l.iSt of Texas); Ohmnberlain, Charleston Mus. Leaflet no. 12, 1939, P• 6 (distribution in South

Oarolina)J Stejneger and Barbour, Check Li.at 0£ North American Amphibians and Reptiles,.4th ed., 19.'.39, P• JO; Carr; Univ. Florida Publs. Biol. Sci., ser. 3, no. 1, 1940, P• 5:3 (distribution in

Florida); Green and Richmond.• Copeia, no. 2, 1940, P• 127 (:f'irst :record from West Virg-l.nia); Petm,. T•nneasee Acad. Sei., vol. 15, no;.,._ 3, 1940, P• :353 (.Alabama); Barbour, Copeia, no. 4, 1941, P• 262 (Kentucky); Freyburger, Proc. 'Permsylvania .Aaad., ·Sci., vol.

15, 1941, PP•· 180-183 (Pennsylvania); Green, The Marshall Review• vol., 4, no. 2,· 1941, P• 36 (additional reaords b-om West Virginia); Wright and Wright, Handbook of frogs and Toads, 1942, PP• 54-55, pl. 9 (general discussion); Stejnegar and Barbour, Cheak List ot North .American Amphibians and Reptiles, 5th ed., 194:3, P• JS; Petrides, Copeia, no. 2, 1944, pp. 122-123 (five years iri oapt1v•

itYt 0 at least s~ years old at time of death")J Anderson, Bull.

. . . \ Chicago Acad. Sci~, vol~ 71 no. S~ 1945,· P• 273 (second.record from Missouri); Conant, soc. Nat. Hist. Delaware, 194.5; P•.) (records from Del.aware, Maryland, V:1.rginia)j Elder, Qopeia~ no. 2, 194.5. P• ·

122 (first, reQord from Illinois); Gier, Copeia, DI?• 1, 1945~ •P• 50

' ' (first record £ram Ohio); Burt, Trans~ Kansa.s,Acad. Soi.; vol. 48, no. 4, 1946, P• 423 (Louisiana.); Obrecht, Copeia, 1:10. 2, 1946, P• 72 (South Carolin~) J' Conant, Reptiles ·c:4'1<:l Amphibians 0£ the Horth•· eastern States, 1947, P• 5, 29 {photograpb); Goin, Cop~ia, no. 4, 1947, p. 275 (prey item ot Heterodon sirnug 1n Florida)J Hoopes, Copeia, no. 2, 1947, P• 138 (in captivity nine years); Livezey and. WrJ.ght, Amer. Midl~ Nat., vol. 37, no. i, 1947, P• 202, fig. 5, eggs described and tigured)J Parker, Proo. Arkansas Acad. Sci.,. vol. 2. 1947, P• 18 (Arkansas); Richmond, Ecology, vol. 28, no. 1, 1947, PP• SJ-6? (larval development and behavior in: Virgini.a); Green, Copeia, no. 1, 1948, p. 65 (saoond report i'rom Ohio); Neill, Herpetologica, vol. 4, no. Ji. 1948, p. · 108 (Georgia); Wright and Wright• Handbook ot Ftogs and Toads ot the 'United stat~s · and Canada, 1949, PP• 12:3-127, pl. 21 (general discussion); Barbour, Amer. Midl. Nat., vol. 44,. no., :3, 19.50, P• 759 (distr.l•

bution in KEtntuok,y) J Brown, An annotated eheek list of the reptiles

anc! :;..mphibians 0£ Texas, 1950, p. 39 ("no valicireoords !rom Texs.G");

' ' Harrison, Pennsylvania Angler, 19.50, p. 22 (diatributionin Pennsylvania)1 Neill, Herpetologioa, vol. 6, 1950, P• 114 (Georgia); 28.

Robertson• Jour. Elisha Mitchell Soi. Soc., vol. 66• no.· 2t 19.50, p. 1,34 (North Carolina}f Spangler,, Ohio Jour •. Sci., vol. 50, no.

6; 19.50, P• 277. (seeond· reoord £rom Ohio)t Viosca11 Louisiana Aead.. Soi .. , Pop,.· Soi •. Bull., no~ 1, 19$0, P• 10 (Lou\.siana); Wilflon and Friddle, .Amero Midl• Nat., vol. 43, no. 1, 1950, P• 168 (West. Virginia); Barbour and Gault, Copeia• ·no. 3~ 1952• P• 192 (natu- i-ru. history 1n Kentucky); Chermook, Univ. Alabama Mus. Pape:• 3.3,

19.52; P• 30 (distribution in Alabama)t Orton, km.er.· Midl. Nat.,

vol. 47, no. 2, 1952,. pl~ .'.l~ i'ig. 1.5 (tad.pole mou.thpal"ts £igured)1 Smith, Copeia, no. 1, 19.52~ '.P• 48 (extension 0£ range in New York); Brandt, Herpetologioa., vol. 9, 195:3, P• 141 (breeding ha.bits in

Georgia); Goin and Goin,' Ecology, vol. 34, no. 2 11 19.5:3, P• 406 (eaology in Florida)J Pearson, JC7U1". Ten.-r1essee Aoad. Sci., vol. 28, 195.'.3, P• 186 (abstraot ot ecological stud.yin Florida)1 Schmidt, Cheek List or North Junerl.oan Amphibians and R1eptiles, 6th ed., 19.53,

p. !58 ( type looality restrloted to Charleston, South Carolina)·; Gosner and Black, Copeiat no. 4• 19.54, pp. 251 ... 254 {larval develop...

ment in New Jer,s&y); QUinby0 Herpetologica, vol. 10, no. 1, 19.54. p. 8 (notes on breeding habits in South Carolina)i Gosner and Black•

Amar. Mtdl .. Nat., vol .. .54, 110. 1, 1955~ pp. 192 ... 203, figs. 1-2, (factors influencing reproductive eyole; distribution in New Jersey);

Pearson, EOl)l., MonograJ)lw, vol,. 25 1 195.5,· PP• 2:;3-267 (population ecology in Florida) •. · Taxonond.g b1stor,v;i Dr. John Edwards Holbrook px-oposed the Genus a,oanh6opus (18,36:8.5) £or the spadef'oot toad he deaoribed aa §.. s0;litslli'.U§• Re was urunmre that one year earlier Harlan (18J,5;10.5) deacribed this amphibian from ·south Carolina under the name of Rana holbroolg.. the o:t G~r (18771194) desQribed. Soa.pW,opu§-. . a:J.bu~. on basis_ . .

twelve t,p$Cimans. collected. by Ootmt . Pom:-tales. . . . ·from 'Key' west, Florida, and subsequently (1884,4.5) reduced §1.JJ;m§ to au.bspeeii'ic rank; he considered " -- . .· . . it oonspeoific· with Soap_pioEB:S; @o,Mtariµ,sp(~.- holg..MOOisi}., Up to recent tilnes ,a. h• alb,yfwas ·regarded as a doubtt'ul subspecies (Deckert• 192h22;_ ca:rr, 1940:,54J Wright and Wright. 1949:129, and others). D11ellman (19.5.51143) ;relegated al.but to ·the qno~ or· a9fi.Ph\PFB.! . holbroolg.. noUng that the coloration ot &1-Jimt was not diagnostic

and that :speoimens .from the mainland o:t Florida afld ice; W'~s:t-0::d:1.d not dif':ter significantly in structure.,

W,agns,QSt. _A Sca"OW,.o;ft,¾§ having pectoral._ glsndsg- interorbital ·elevation not strongly developed; only slightly ~onvex;_ tympanUlll- -

and parotoid distinct, the £orm1.u:- 4. 1 mm. to 5 .. 9 mm. in diameter (av. 5.1 mm.),. _the latter round; sides or frontoparl.etal bones d1vergent to a point. near poster:Loi" 'bord.al'-_ o:t orbits, converging

strongly just po$terior to this point (pl. I,. fig. J)J £oramen

magn'lll11 bxtoader than longJ few or no blaQk spines on head.

)Cmat-;on: No consistent var.i.ation that oould be oe>rreJ.ated

with geographical distribution was noted, and no int&rgrades are

known between J-Ao),bl:oold. and !ffirte;ga. The latter bas been :regarded .

by maey as a subspecies of SogphiO"f?UEi holbroolsi• As evident upon

exam1ning the map, figure 1. the geographic :ranges ot the ttto kinds

do not overlap. On4" 1n the state of Arkansae are both to be found.

and. a wide gap exists be¢.ween the known ranges of eaoh in that State. Qenergl dist;1;butiona From Massaehusetts southwester~ to south• eastern Misaourl, northeastern At-kansas southward into eastern Louisiana. Although Wright and Wright (9.2. ~.) and Schmidt (1953;,58) place extreme aou.theastern Texas within the range. of the solitacy spadef'oot,. Brown (19S0sJ9) remarks that there are no valid teoords 1"rom Texas £or this speoies. Elder (19451122) recently added this speoiea to the fauna or IllinoisJ its presence there was conti..'l'"llled b:r Smith (1948t2). Scaphiopus holm;:ooJa 1-1as discovered reoentl;:r in Ohio by Gier (1945:.50) J Green

(1948:65) and Spangl.er (19.501277) con:tir.med the occurrence ot the species in that State. Fragmentary fossil material from the Miocene of Florida is referred to this speoiea by Auttenberg (19.56t6) ..

§Paoimenm exa;nin§ch ·S7; &-Jsmisa~: Car::iway,. Craighead Ot>. (2 KU); Imboden, Lawrenae Co. {1 KU); Florida~ OaineavUle (1.5 KU), 4 mi. S of·

Gainesville,. Alachua Co. (6 KU); XentuokY: Jopfa Ridge (17 KU) 11 Mammoth

Cave Ridge, Edmonson Oo. (2 KO')J M3:tYl4Bdi Massay, Kent Co. (4 OJS); Ma;1sachusl.\tt§t, Middleboro, Plymouth oo. (1 KO); ~rginia: 2 mi. N of Kenova, Wayne Co .. (9 KU).

SoapbiOpwJ huz;:ter;l st.reeker

(pl6 I, fig. 2J pl. II, fig. 3; fig. 1) Boa.phiopus bglbroold. .. Strecker, Proc. Biol. soch waabin$ton, vol. 2:1.,

1908,. P• 52 (Te:xa.s)J Bragg, Amer. Mid.lo Nat., vol. 24, no. 21 1940, P• ,32 (eoologioal distribution in Cleveland. Co., Oklahoma.).

Scaphiopya hurtetli streoker, Proo. Biol.. soo. l-Tasbington9 vol. 23, July 23,

1910, P• 116, pl. II. Typa locality: , :3½ miles East of .WaooJ Motennan County, Texas; Stejneger and Barbour, Check List o:r North -· 92

0--=:::;===~300 • specimens el!Offllned . scale of miles

o other spec·1mens

92 78

Fig. 1. American Amphibians and. Reptiles1 1917; P• 26t' . Nieden•. Das. Tierreich, Lief. 46J Anura I, 192.3; P• 46t Stejneger ancl Barbour, Oheak List of North . .Amerioan.Ampbibians and Reptiles, 2nd ed., 192.3,.p. 2,3; St.reeker. Bull. Baylor Univ., vol. 38, no • .'.h 193.5, p .. ,, (Tsxas)j Smith,

Herpetologica, vol. 1, 19.37. PP• 104.,.108, pl•. 11 (mu;:tetj. is distinot from holb1:001d.;· akull figured; tint record :from.Arkansas); stejnegor and Barbour, Cheak Li.St of North .American Amph:1.bians and Reptilos, 4th ed., 1939, P• JO; Tanner, Great Bru!dn Nat., vol. 1, no. 1, 19.39, PP• a... 9, pl., 1, figs. 3-41 pl. 2, fig. :3 (critical taxonomic study);

Bragg, Turtox News, vol. 20 9 no. 1, 1942s P• 12 (breeding behavi.or);

Bragg, Soieno$, vol. 95 1 1942, Po 194 (behavior in Oklahomah Bragg, Proo. Oklahoma Acad. Sci., vol. 22, 1942, P• 16 (Oklahoma); Bragg,

Anat. Record, vol. 84, 1942 11 P• 506 (life history in Oklahoma);

Bragg, Turtox News, vol.• 201 no .. 11, 1942, p.1,54 (Oklahoma); Bragg and Smith, Great Basin Nat., vol .. J., no. 2. 1942, PP• ~39, 47, 49. (breeding behavior in Oklahoma); stejneger and Barbour• Cheak

List 0£ North American A.1T1phib1ans and Reptiles, 5th eda, 194:3, p ..

;8; Bragg, Amer. Nat •• vol. '78, 1944, pp • .517-53.3 (distribu.tion in Oklahoma.J critical. study of ha.bits; SUllllJW7 ot knowledge of species) J Bragg., Copeia, no. 4, 1944, pp., 230-241• figs. 1.. 2 (breeding habits, eggs, tadpoles discuseed; tadpole, egg figured); Bragg, Amer. Nat .. , vol. 79, 1945, pp"' .52-72 (breeding patterns, pbylogeny)J Bragg,,

ppe and Wasmarm COolleotor, vol., 6, no. :3, 194.5, ·, ' 69-78.. . (breeding tadpole behavio:r in Oklahoma)J Bragg• Prooe Oklshoma A9ado Sci., vol. 2.5, 1945, P• 27 (puzzles in life hiStory); Braggp. Proe •. Okla... homa Acad. Soi., vol. 26, 1946. P• 19 (aggregational. phen~eua in tadpoles)J Bragg, Wasmann Collector, vol. ?, no. 2, 1947, P• 66 (tadpole aggregat:1.ons)J Bragg, Proo. Oklahoma Acad.. Sci., vol. 27, 1947, P• 61 (tadpole behavior); Bragg, Anat. B.eoord, vol. 101, 1948~ p., 706 (.feeding behavior of tadpoles); Braggi Proo. Okla- homa. Aoad. s01., vol. 29, 1948, p. · 11 (dermal saoretions by tamales . . . . suggests sexual di££erence)J Bragg, Wasmann Collector, vol. 7, 1948, P• 66 (social aggregation in tadpoles}; Bragg, Proc. Oklahoma. Acad. · Sci., vol. 30, 1949, PP• 18-.:1.9 (describes egg masses); Burger, Smi·th, and Smith, Jour. Tennessee Aoad. Soi., vol. 14, no. 3, 1949, P• 1:;o (s~cond report from Arkansas); Bragg, in Researches on the Amphibia 0£ Oklahoma, 1950, PP• 27, 3.5, 78, 10:1.. (distribution, breeding habits, adaptations of survival vaJ.ue in Oklahoma diaaus:,ed)J Bragg, Proo. Oklahoma. Aoa.d. Sci.• vol. :31, 19.50, PP• 26-27 (mass moveioorrt. at metamol"phO$is)J Bragg, Wasmann Jour. Biol., vol. 8, no. 2; 19.50, PP• 221.228 (social aggregations of tadpoles); Bragg, Proo! Okla- homa. Aoad. Sei., vol. J1, 1950, P• Z14 (extension of kncmn range in

Arkansas) J Brown, An annotawd check list of the reptilea and amphibians or Texas• 1950• p • .39 (d1strlbuM.on in ',t'exas)J Peterson, - Arner. Midl. Nat., vol. 43, no. 1, 1950, P• 158 (Texas); Reese and Firsobein,. Trans. Kansas Aoad. Soi., vol. s,, no. 1, 1950, P• 49 ('l'exas)J Bragg and Dundee,. Pl'OC. ·Oklahoma Acad. Sci., vol. :,o,

19.51, P• 20 (Oklahoma); Bragg, Wasmann Jouro Bi<>l.;, vol. 10, l'.D. 2, 1952, p .. 249 (Oklahoma) 1 Smith and Sanders, Texas Jour. soi., vol. 4, no. 2, 1952,. P• 209 (additional records from. Texas); Axtell and Wasserman, Herpatologica, vol. 9, 19.53, P• 2 (records

trom southern Texas)I Bonn alld MoOarley• Texas Journ. Soi., vol. 5, no. 4, 1953, p. 467 (Oklahoma); Bragg, Herpetologioa, vol. 10,

no. 21 1954, PP• 97.. 102 (tadpole behavior in Oklahoma.)J Bresl.er

and Bragg, Copeia, no. 4, 1954, P• 256 {variation in rows o£ labial teeth of tadpoles); Blair, Texas Jo'\U". Sci., vol. 7, no. 2, 1955# P• 185, fig. l (sonogram of call). :a2ap~ows holbrooki hol.broolq, ortenbu.rger, Copeia, no. 163, 1927e P• 46 (Oklahoma); stejneger and Barbour, Check t:1.st o£ North .American Amphibians and Reptiles, :3rd ed., 1933, P• 26 ~.); Smith and .Leonard, Amer. Mid1. Nat •• vol. 15, 1934, P• 191 {Okla-

boma.)1 Black and Dalllnger1 oooas. Papers Univ. Arkansas Mus., no. 2, 1938, P• 13 (12art.).

Scapltj.opup lJ2lbt90k1 hur:teti• Wright and Wright, Handbook of Frogs and

Toads, 193.3, PP• 44-45 (generu disoussion); Wright and Wright,

Trans. Texas Acad. Soi., vol. 21, 19,381 p. 18 (distribu.tion in

Texas); Wright and Wright, Handbook of Frogs and Toads, 1942,

pp~ S7 (general discussion); tivezey and Wright, Amer. Mid.le Nat., vol. 37, 1947, pp .. 193,203, £1g., 6 (eggs desorlbed and figured); Blair, Copeia, no. 1; 1949, P• 72 (development, in Texas); Wright

and Wright, Handbook of Frogs and Toads of the United States and Canada, 1949, PP• 130.134, pl. 23 (general discussion); Jameson. Copeia, no. 1, 1950, P• 61 (Texas); Schmidt, Check List ot North American !Amphibians and Reptiles., 6th ed., 19.53, P• .58. Taxono¢c hi§tOt:(I This speoies has had a varied history. Certain authorities regard 1t as a subspecies of holbxyoki.. others place it into the synonymy of holbrgold,• and still others maintain it is a distinct species. St.ejneger and. Barbour treated hµrter.tas a distinct Speci~s in the i"irst and seoondedit1ons of their check list (1917• 1923)• ,In the third edition of the oheak list (1933t2J) it was regarded as a

__ , .. ,, synonym o:t Scapbiopus ·. holbroold. bolqrgolg,. Smith (1937:104). on the basis ot the stru.etu:re o:r the akull and certain external. oha.racters,·conoluded. that hurter!. was•specifioall:y distinat from holbrooki. Tanner (19.39:9) agreed with the views ot

Smith. &--agg (194.5:69) reported on the breeding behavior• metamorphosis, and larva 0£ .· h~r+ and noted that these aJ.1 .. differ in ·detail from that of b~oki as described by Ball (1936:.'.351) and others. The results ot Bragg's atudy thus supported tb.a views o£·Smi.th and Tanner. In subsequant cheok lists ot Stejneger and. Barbour (1939, 194'.3), hurten was·given.f'ull speoi.t':\.c rank .. But in the Il10St :recent check listi Schmidt (195,:58) reduces it to subspe(d.fio rank. In nw discussion ot holbrookt I note that no inte:rgrades ex1.st between it and h:ur;teri .. and that their raopactive ranges, a.s lmown, do not overlap. Until intergrada.tion between the t1ro is demonstrated, the logioal course is to treat both as full species. Diagnosis: A L,caphiopU§ ·with peotoral glands, intero:rbital ·· elevation strongly devoloped; tympanum and parotoid distinct, the :f'ormer S trml• to 7 .2 umi. in diamater (av. 6.4 mm.);. the latter elongate; sides o:t :f.'rontoparietal bones are parallel or nearly so, converging slightly at a point near posterior border of orbit (pl, I, fig. 2); fora.men magnum as long as broad; numerous small; black spines on head, reduoad 1n number between and anterior to eyes.

yarl.a.tiop,s · Nearly all of the speed.mens ax.a.mined dif'tered 1ri some' respeots from the description ot Strecker (1910:116-117) and the aooounts of Wright and Wright (1949:131). The parotoid appears more oval than round and the tympanum i~ larger than in specimens ot ho;trbrooki ot like size. strecker (im,• pit.) remarks ttparotoida nearly ·round••• tympanum distinot but rather smaller than in holbrook;\.. (In type hardl,y~ more than half the diameter of the parotoid.)tt I am or the opinion that

the two _specimens which served as the basis £or the description of )J.m.j:.eri were not typical 9£ the populat:i.ono This seems to be substantiated by the fact that Strecker £ailed to mention the interorbital elevation. a.most distinct character in typical hprj;eri. Distrigut:t6n:. Eastern Oklahoma, south-central and eastern Texas,

and western Arkansas. Axtell and Wasserman (19.53s2} report the presence

o:t Scaphiopus hurteri from near Rio Grande City 1n extreme southeastern Texas. It probabq occurs in Mexico although as yet unrecorded from there. In Oklahoma this species.inhabits woodl.and-and savanna areas, aocord- ing to Bragg (19441.528)., In Texas ,a. hurlaV, also- enters more mric veg~tation types such as mesquite-chaparral, judging lrom the distribution of locali.ty·:reaords in.that State.

$:pegixneng, ,xa.minedi 176; ,Arka.ssaia 2 m:1. E ot Ozark, Franklin Co.,

(1 KU); O,kla.qoma: Lexington, Olevala.nd Co,. (2 EHT-ID.fS)J Te:xaa2 Benton (2 I{{J), cytle, Atascosa Co. (4 KU); Somarsat, Bexar oo. (169 KU).

Soaph19pu.§. couohi Baird

(p. I• fig. 11 pl. II, figs. 1, 2; pl. III; £ig. 2) Soaphiopua pouchi Baird, Proq. Acad. Nat. Soi. Pb:Uadelpbia, vol. 7, / April, 1854, P• 62. Type looalit:n Rio Nazae, Coahuila, and Matamoros, Tamaulipas, Mexico; Baird, United States ,Mexican

Boundary Survey under the order ot Lieut. Col. w. H. Emoey-1 Reptiles of the _Boundary; vol. 2, 1859, P• ·28, pl. :35• figs. 1•6

("Lower Rio Granda0 ); Oop&, Proo. Aoad. Nat. sc1 •• -Philadelphia, 186:3, P• ,52 (Baja Ca.11£ornia}; Copa, Jour. Aca.d. Nat. Scio Philadelphia, vol. 6. ser. 2, 1866, p. Bia (liated}r Cop&; Bull. u. s. Nat. Mus., no. 1,1875, P• :32 (listed); Brocobi,·Mission

. / ·. . / scientifique au Me.xique et dans l*Amarique Centrale, Etudes sur . 1es Batraoiens, livr. 1, 1881, p. 26 ("Tamaulipas• Rio. Orande")J Boul.enger, Catalogue of the Batrachia Sali.entia ilr th.a- British

. Mus8illl1, 2nd ed.., 1882, p. 4'4J Yarrot.r, Bul.1. u.. s. Nat. Mus.,

no. 24, 1883, P• 177 (p~.) J Garman, Bull. Esse.."'t :rnst. • vol-, 16, 1884, P• 46 (~.H Belding, West Amar. Soi., vol • .:,, no. 2.4, 1887, P• 99 (Baja California) J Garman; Bull.. Essex Inst., vo1. 19, / . 1887, P• 20 (Coahu.1.J.a, San Luis Potosi);Cope, Bull~ u. s. Nat.

l'ms., no. :34, 1889,P• :301 11 .fig. 75s pl. 68 11 fig. 2; van Denburgh, P:roo. Oa.lliorn.\a Aoad. Soi., vol. 5, ~er. 2, 1896,- P• 556 (Baja

vol,. 111 Oal1:rornia);. Mocqua:rd, Uouv. • .Aroh.. Mus. Hist. Nat.; ser. 4,

1899, P• .'.}42, pl. 13 11 fig. 6 (:f'oralimb figu.red.)3 Ounther, Bio1ogia

CentraJ.i... ,Amerioana, Reptilia and Batraohia., 1901,- P• 259 (Baja. O~lti"omia., Texas, Mexioo, Presidio. Tampicop Tmnaulipas)J Diakerson•

The Frog Book, 1906, P• 57, pl., 8, col. pl. 2 (general diacussion);

Ruthven, Bull. Amer. Mu.a, .Nat. 1Iiste, vo1., 23, 19071 p. 503 (Arizona);

Strecker, Proc. Biol. Soe. Washington, vol. 21 1 1908 11 p. 52. (Texaa)J Strecker, Proo. 81.ol. soa. washix1gton, vol. 21, 1908, P• 60 (additional 1ocall.t1ea in Texas); st.reoker, Proo. Soe. Washington,

vol. 21 11 1908, PP• 199-206 (distribution, notes on Ute history); Streaker, Baylor Univ.- Bull., vol. 12, no.1, 1909, p .. 9 (Texas); Strecker, Trans. Aead. Sci. St. Louis, vol. 19, 1910, p. 81 (Waco, Texas); Stone, Proc. Acad.. Nat. Soi. Philadelphia, 1911,

P• 22;3; Van Denburgh and Slavin, Proo. Oa.lif'o:rnia, Aoad. Sci., ser. 4, vol. :,, 1913t P• ;395 (A1"i.zona.)J Stejneger and Barbour, Check ti.st or North luner1.can .Amphibians and Reptiles, 1917, P• 25 (distribution) I Van Denburgh and. Slevin, Pron. Oal.i£ornia Acado Sci., ser. 4i vo1. 11, 1921,. P• S3 (Baja California}; Nelson, Mem. Nat. Acad. Soi,., vol. 11, no. 1, 1922, P• 113 (Baja Calitornia); Schmidt, Bull. Amer. Mus. Nat. Hist .. , vol. 46, 1922, P• 6,3, (su:mmar.v ot distribution in Baja Calii"ornia); Nieden, Das Tier:reioh~ Liet. 46; Anura I, 1923, p .. 47; stejnegei- and Barbour, Cbeok !,1st ot North American .Amphibians and Reptiles, 2nd ed.,

19231 P• 22 (range); Ortenburger, Pro"• Oklahoma Acad. Sci., vol. 4, 1924, P• 19, pls. 2... 3 (habits, eggs; Arizona.); Van Den• burgh, Proo. California .A.cad,. Sci., ser. 4l> vol. 13, 1924, p. 195 (Net-t Mexico); Ortenburger and ort.enburger, Proo. Oklahoma Aoad. Sci., vol. 6, 1926, P• 101 (habits in Arizona.); strecker, Contrib .. Baylor Univ. Mus., no. s. 1926• p. 11 (lif'e history, Texas)t Slevin, Occas. Papers Oalifor:rda Acade Sci., no. 16, 1928, P• 87 (general discussion)J Strecker, Contrib. Baylor Mus., no. 1.5, 1928, P• 7 (Texas)J Wright, Proo., u. s. Nat. MUB., vo1. 74, 1929,

PP• 4, 20 1 pls. l, 6 (tadpol~s described and figured)& Noble, The Biology o:t tha Amphibia, 1931; PP• 4oo, 488, figs. 1.35, 1.5.5 (vocal pouch and vertebral column figured); Kellogg, Bull. u. s. Nat. Mus., no. 160, 19.'.32, pp., 20-21 (description, di$tr1.buti.on in Mexico); King, Oopeia, no. 4, 1932,. P• 17.5 (Arizona); Linsdale,

Univ. Oalli'ornia Publs. Zool., vol. :38 11 no., 6, 1932, p .. .).50 (Baja. Cal.1£ornia)1 Allen, Oooas. Papers Mus. -Zool .. Univ. Michigan, no. 2.59t 1933• p. 2 (first record .from Sonora; Metlco); Stejneger and Barbour• Check Li.st of North .Amer1oan·Antpb1bians and Reptiles, 3rd ed.;

1933, P• 26; Wright' and 'Wright, Handbook of Frogs and Toads, :19:33,

PP• 38-39, pl. 6 (general discussion); Smith and. Leonard,, Amer. Midl. Na.t.~ vol. 15• 19:34, p. 190 (Oklahomah Chao'Liu, Proc. Boston Soc, Nat. Hist., vol. 41, no~ .3, 19.)5, P• ;O (VQoal sac desor.tbed)J Dunn, Proo • .Ad.ad .. Nat~ Soi., Philadel.phi.a, vol. 88,

1936, P• 471 · (San Luis Potoa£: !i!e:d.coh Gloyd, Bull. Chicago Aoad. Sctl.., vol. 5, no. S, 19:37, P• 10:3 (Arizona}; Litt.le and Keller, Cope:ta, no. 4, 19:;7, P• 218 (Nati Mexico); T8¥lor, Univ. Kansas Soi. Bul.l., vol. 24, no. 19, 1938 (19.36); P• -478 (Sonora, Mexico)J Ta;y'lor, Unive .Kansas Soi. Bull., vol. 24, no. 20; 1938 (1936), P• .$08, pl. 44 (.ti.rat reco~ from Sinal.oa, Mexico); wright and wright, Trans. Texas Aoad. Soi., vol. 21; 19:38; P• 17 (distribution :l.n Texas); Stejneger and- Barbour, Oheck List of -North

Amer-loan Amphibians andRepti1es1 4th ed.,- 1939, p. 29J Tanner. Great Basin Nat •• vo1. 1, no. -1. 19.39, P• 10, pl. 1. figs. S..6t pl. 3 (critical taxonomic stud;Y); Little• Cope:1.a, no. 4. 1940• P• 262 (habits in .Ar:lzona)J Bragg, Copeia, no. 1, 1941, p. S2

(Oklahoma records); Bragg and Smith, Great Basin Nat •• vol. 3, no. 2, 1942, PP• -3a ...:;9 (breeding behavior· in Oklahoma) J Bragg, Turtox Nar..rs. vol. 20. no., 11t 1942, P• 1.54 (Oklahoma)J Wright and Wright• E:andbook of Frogs and Toads, 1942, PP• 46-47, pl-. 6 - (general diecussion);_Arnold,, Cope1a, no. 2, 194.3, P• 128 (behavior in Arizona); Kauffeld• .Alner. Midl. Nat.,, vole 29, no. 2, 1943,

P• 342 (Arlzona)J Stejneger and Barbour, Check List of North

.Amerl.can Amphibians and Reptiles. 5th ed,, 194.3,, pe 37; Bragg, Amer. Nat., vol. 78, 194411 PP• .517•53:3 (SUlll1l!B1"Y of knowledge ot life history; distribution in Oklahoma)j Schmidt and OWens, Field Mu$. Nat. Hist., Zool. aer., vol. 29, no. 6, 1944, P• 98 (records tromnorthern Coahuila, Mex:i.00)3 Bogert and Oliver,

Bull. Amer. Mlls./ Mat. Hist., vol. 83, no. 6, 1945, P• 339 (Sonora• Mexico) J Bragg, Amer. Nat., vol. 79, 1945, PP• ,52-72 (bl"eeding patterns, phylogeny); Taylor and Smith;' Proo;,; u.. s. Nat. Mus., vol. ·9.5, 194.5; P• .5.51 (Chihuahua and Tamaulipas, Mexico); Bl.air;

Copeia, no .. 11 1947, p.· 67 (breeding behavior in Coahuila, Mexl.oo);

Bragg, Proc. Oklahoma A.cad. sQi., vol .. 27 1 1947, P• 61 (tadpole bohavior)J Livezey and'W'righh Amer. Midl., Mat., "Vol. Yl, 1947• ,,

PP• 192, 202• fig. ) {eggs described and figuredH Bl•a.gg 0 Proc.

Oklahoma Ac:ad.. SO'i~ 0 vol. 29, 1948, p. 11 ( 11derrnal secretions by females suggests sexual di.tferencen); Bragg and nundee,. Proo.

Oklahoma Aoad. Sci., voi. 29 9 1948 11 P• 2.5 (Oklahoma.)J Smith and Taylor, Bull. u. s. Nat& Mus., no .. 194, 1948; p.; ).5 (distribution in MexJ.eo sumznarized)J Burger and Hensl$Y& m1t. Hist .. ?1isoo 1 no11 JS,

1949, P• 2 (Sonora, Mexico); Conant a.nd Hudson, Rerpetologtca, vol. S, 1949, P• ) (3 yt!tars, 4 months in ¢aptivity); Jameson and Flury, Texas Jour. Soi., vol. 1, no~ 2si 1949, P• 59 (Texas)J Maidonado-Koerdoll, Rev. Soo. Mexioana Hist. Nat., vol.. 10. nos. 1-4, 1949, P• 247 (new locality- re

(distribution and 'breeding habits in Oklahoma); Brown, A.ti Annotated Cbeoli: JA.. st of tho Reptiles and Amphibians of Texas,· 19.50• ·p. :38 (distribution in Texas) J· Firschein, Herpt:,to1ogica, voJ.. 6, 1950.

p.: 75 (Cbihuah~, 1'...ex:teo)3 Reese and ~sobe:iri, Trans. Kansas

Aoad. s01 .. , vol.5.3, no. 1, 1950,·p. 49 {reeords from Texas and Tmnaulipas; Mexico); Smith and T~lor, Univ. Kansas sci. BuJ.J•• , vol. ,:,::3, no. 8, 1950, P•· 345 (type loQality restnated to Matamoros,

Tama.ulipaa, Me:.rl.co}; Bragg and Dundee. Proo •. Oklahoma Acad. sc1 •• vo1. Jo, 1951, p. 20 (Oklahonte. records); stebb:i.na, Amphibians of western North .Amerioa, 1951,. pp. 197-201, pls. 14• 2.5, 27• 28, .55 {general disoussion) J Teyl.or, Univ. Kansa.e Soi. Bull., vol. '4, / n<>. 13, 1952, P• 797 (San Luis Potosi, Mex:1.ao)J Axtell and Wasuer-

tna11, Herpetologica0 voi.: 9, 19.53, P,• 2 (Tex.aS)J Bbad:url., .Proo ... Zool. soc~ Bengal, vol. 6, 195:3, P• 18, fig. 2, pl. J, £iga. 14-16 (u;:-ogenital system discussed an

Taylor, Un1.v. Kansas Sci. Bull.1 vol. 35, no. 2, 19.53, P• 1590 (additionaJ. localities in San !;uis Potos{. Mexico); Stebbins, Amphibians and Reptiles of western North .Amerl.ca, 1954, PP• 9!~, 141, pls. 11• 17, fig. 13 (general aaeount; distribution);Blair• Texas Jour. Sci., vo1. 7, no. 2, 19.55• p. 185~ fig. 1 (sonogram of can); Duellr.ian, O~oas. Papers Mus. ZOol. Univ. Michigan, no • .569• 19.5.5• P• 2 (A'r':l.zona); Walters, Copeia, no. 3, 195.5, PP• 2,52-2.53 (.Arizona) J Zweil'el and ?-l'orrls, Amer. Midl. Nat •• vol.- ,54, no. 1, 19.5.5, P• 230

(:; looal.itiee in Sonora); Jameson, Cppeia, no •. 1 11 19.56, p .. .55 (rocoveey o! marked speoimens in Texas). 41.

§.(eaplrl,opg,s) varj.J.Yi Oope 11 Proo. Acad. Nat. sci. Philadelphia., March,

186:3, 'P• .52. Type. locality: Cape San l,uoas, Baja Californiai Oope, Bull. U. S~ Nat. Mus., no. 17, 1880~ P• 29 (Texas); Brocchi,

Mission Sci. Me:xi6ana, Batrach., pa.rt .3~ seot. 2 1 1881, P• 27 ,: (listed).

§(P§Lphio;qus) rmctif'renis Cope, Proo. Acad. Nat. Sci. Philadelphia, March; 186,;, p. 5:,. Ty:pe local.it;n Tam.aulipas and. Coahuila, Mexico i Broochi• Mission Soientifique au. Mexique et dans l'Amerique Centrale; Etudes su:r les Batraoiens, livr. 1,, 1881, p. 27 (listed)J Boulenger, Cat. Batr. Sal. Brit. Mus., 2nd ed., 1882, P• 4J5 (listed}; Garman, Bull. Essex Inst., vol. 16, 1884, p. 46 (listed); Gunther, Biologia. Centra.li.•Amerioana, Reptilia and Batraohia, 1901, P• 2.59 (di.tair.lbution)J Nied.en, Das ~1erreich,

Lief. 46; .Amn-a I, 192:3, P• 471 Smith and Taylor, Univ. Kansas Sci. Bull., vol. 3.'.3, no. 8, 19.50, P• 328 (type looality restrioted to Rio Nazas, Coahuila, Mexico).

Scapbiopu.a ooµoh1 xarJ.AA Cope,, Proc • .Acad. Nat. sci. Phil.adelpbia. 1866 11

P• ;31) (listed); Garman, Bull. Essex Inst., vol. 16 9 no .. 1, 1884, P• 46; Cope, Bull. u. s. Na.t. Mus., no. 32, 1887, P• 12 (listed); Cope, Bull. U.. s. Nat. }fus.; no. 34, 1889, tig. 75• Saa:pW.oWf yarius v¢us Cope, Bull. U. s. Nat. Mus., no. 1, 187.5, P• 31 (listed)J Yarrow, Bull. u. s. Nat. Mus., no. 24, 188:3, P• 177 (listed). Scaphiopu.s variup rect1£,renis Cope, Bull. u. s. Nat. Mus., no. 1, 1875, P• 32 {listed) I Yarrow, Bull. u. s. Nat. Mus., no. 24, 188.3, P• 177 (listed). ?~ :J,atioaps Cope, •. Nat., vol. 27,, Feb. 189.3,.P• 15.51 1 tig.

Type looalityt 0 between Seymour, in northwest Texas south ot the Red River,. and .Austin." } ~- · ?;3gapMopu.s latieepa., Ni.eden, Da:t Tierreich,. Lief'. 46J Anura· I, 1923,

a.e_m,hi2m1a oouql'.4 ogp.ehi., Smith and Sanders, Tex.as Jour. Sci •• vol. 4,

no •. 2, 19.52, P• 209 (Texas); Smith and Darling, H$rpetologioa, vol. 8, no. :3, 1952, P• 81 (three looalities in Tamaulipas, Mexico). Sgaphigpus couo!q ractifrenig, Smith and Sande~s, Texas Jou:r. sci.,

vol. •. 4, no •. 2, 19521 p. 209 (d1$tribu.tion in Texas)J Langabartel

and Smith, Herpetologica, vol. 10, no. 21 19.54, P• 12.5 (three looal.1ties in Sonora, Mexico). aiaxonomi,o ·h,\s~• Baird ( 1854:62) described SoQpbiopus couch;! from. thrGe speoimens collected. by Ueu.t Danus N.· douoh 1n· 18.53 from two localities in Me:x:too (R(o Nazas1 Coahuil.a, ·1 specinien; Matamoros, Tamaulipae, 2 specimens). In bis original desoription Baird neglected to cite the catalog mmibers of the: oot;ypes.. Cope, however, revised . . ' the Genus in 186:3• and remove ,a. ;:eotifrenis. In the same paper Cope (1863:.52) OitedU. S, N. M. no.

:3713 from Tamaulipas a$ the basis tor the d1.agnoais of§.•· eouohl.. Aooording to Kellogg (1932bt21) none of the types is extant. He adds that one cot;rpe of ;g:ggt2;frem.a (U. S. N. M. no. 371.5) from Tamaulipas is marked in the record book of the division of reptiles as ha.Ving been destroyedJ the.others cannot be located. 4J.

Oope (1889:.301) placed §

The type locality 0£ aca.phioPWf gouc!Y, has been t-estr.lcted to. Matamoros, Tamaullpas, Mexico by Smith and Tay1or (1950::,45). '•' ) In the sixty-three years following Oopo•s description (189J:1.55) of·~ laticims i"rom wests~. Texas, oncy a. single author (Ni.eden, 1923:49, treating it as a tull speoi8s) to·~ knowledge• has oited thia species •. All .Americam workers, strangel;r enough, have t,tompletely ignored it. I here· treat it as a· questionable synorzym of ~§Rlp.opus couobi on the :following grounds. In the original description Qope (sm• ~-) remarks "this interesting speoies approxi.matas more Q1osely the genus scapbiopus than the lmown speoies or Spea. Its colgr (italics mine) and the posit1on·ot its vomerine teeth are more like those or qcaphioPM oouchi ••• n.

The name J&Upe:g§ is not ref'era.bl&i in nw opinion to SpaphioJ2U! lpu:terl, also occurring in Texas• $cein the original description Cope (gn._Q1.) notes "there are no pale longitudinal stripes,tt and "tympanUlll conoealed"J no mention is ma.de 0£ a post. orbital boss or'pectoral glands and the t:rpe locality (•between

Seymour 1n northwest Texas south of the Red Riwr, and .Austin.") is outsido .0£ the known rah;ie ot ~. hµrt§r:t. On the strength ot the above evidence~. b,u;:teri is seemingly precluded from consideration. . . Dr. Doria M. Coohran informs me (in litt.) that- the type speoiman 44.

of la;tioaps is not in the U•. S. National M\ls$Ull19 nor was 1 t in• oluded in the list (prepared by the late E:. R. Durin) 0£ typo spacimGns in the museum of th~ Philadelphia .Academy of $ciences. pia.gno§iea A Scaphib';rnis lacld.t)g pectoral. glandsJ no interorbital elevation; tympanum distinct or o<,oasion~ partially concea1ed by eypertroplzy' ot the parotoid• w~oh is variable -in shape. and usual.zy small;. two lal-ge metaca.rpaJ. tubercles on palJ:nJ dorsal

yarj.e.tionr The nmne rea:Yfr~mi§ was revived by Smith and Sanders (19.52,209) to represent. the western (west of about 101°L west) subspecies of Sga12h19pyf_2oucn:;,. the· eastern subspeoies being the nominal :torm. They report that ,1 • .t• t!ot:trrenis differs from~ • .!!•·· souobi "in having a larger adult size and a moi-e unii"ormly X'etioulatod pattern (when present).11 In addition they add that "indiViduals 0£ Jl• cow;:;h! are smaller. darker, and their patterns tend to possess a preponderance ot dark pigment which usually vaguely outlines a pair ot sinuous paravert.ebral light lines. n Oonai,de~abla variation in color pattern is exhibited by spacimens from the western part or the range .,of ~oapM!?Im@. couohi t,gti:rrem;i}.

(as defined by Smith and sanders). •Individuals from the eastern part of the range of' this subspecies (tvestern Texas, .New Mexico, and southeastern

Arizona) generaf.4, possess numerous small bla.ok tubercles {warts) over the .faintly eVident dorsalretioul.ated oolor pattern (see pl. n, fig. 2), g1ving them the appearanoa o£ being sprinkled with black pepper. The spade reaches the posterior margin of the eye. Specimens £rom

Coabuila, Durango, Tamaulipas, and Zaoatecas agree in most Nspects with the precedl;t1g1 but tend to rea.~h a larger $ize (pl. II, fig. 1).

A series 0£ sixteen individuals from central za:oataoas vary in snout-vent length :f'rom 62 :mm. to 71 nnn. (average, 66 mm.). ~- .£• r,.,stirrenis; (of Smith and Sanderi,., .lg,g_., cit.) from south- central A1":tzona. (KO nos. 14536 ..37), western Sonora (EBT-mrs nos. 3812,

3814--1.5, i'rom Ouaymas). ~a (KU noa. 29888-89, from Rosarl.01 KU no., 37826 frgn Ins Moobis, pl. II, fig,. 2), and extreme southern Bijja California (Sl'AN nos. 1.5665 ...79 trom La ru.vara.) are smaller, possess a more diatino"" .finely reticulated dark pattern, and have a shorter hind leg, the spade saa.roe4' reaching the tympan\Ull. Tb.Q peppsrllke spots are absent. However, in Specimens trom ~eal'" Llano (STAN nos. 1_0298, 10;301, ·10306 ...7) and Cocor.tt (STAN no. 10300), Sonora, and La

Paz, Baja California (srAN nos. 1.56.58-6.3). the reticulated. pa.tteJ:"n is absent or oncy faintly discernible• the ground color is light or dark,· and the 4ol"sum is covered with pepper:l.ike spots. One specimen (STAll no. 15664, trom La. Pas, Baja Qalifornia) dif'fers £rom others from that locality 1n having the dorsal reticulation dietinct, but agrees with them in having a pa.la ground color. These spad.efoots all agree in having the skin of the head coossi:f'ied with the skull• the tympanum. small bu.t distinct and the parotoid small,: often 1nd.1stinot. Taylor (1938:,508) reports that in nineteen speoira£i>ns from near

Mazatlin, Sinaloa the skin ot the oooi.plta.l and interorbit.al region is smost completely involtred in the ossifioation of the skull, despite the .faot that. none appear to be £ull grown. Mo:re recently Zweif'el and 46.

Norris (19.5St2,31) report ape~ trom south of Rerniosillo, Sonora. that c:liffei- .from northern examples at Gu:troooba• Sonora, in having a smallez- $lie• a lighter ground co:toz-. and the skin o£ the lwa.d ooossified with the cranium even in the S!11alleet :1.ndividual. (41 mm. snout-vent length). The smallest speoinien available to me (STAN no.

15675 1 tx-om. I.a Rivera, Baja Calitonrl..a) is a female. It measures 34 mm. ono11t-vent. and the skin o:t the head is ooossi:tied with the skull.

No eggs a.re pzi.,sent in the body cavity. A $omev,rha.t larger £emal.e 9 52 mm. snotrb-vent, tram the same locality contained. eggs and. also had the:"sld.n o:t the head involved in the essitioat:,..on of the skull.; Certain specimens tram. Arizona, Baja aalitol"llia. sdziora, and

' ' Sinaloa, aa noted above, can be ~erred to· ,a. R.• .pouchl::~'(tht;t presumed Ellastern eubapeQies} op. the basis of' dorsal ool<:>r pattern !!ml size. others a.N referable to §. • .2.• regti.fr;9,m,a ju.dgi.ng !:rem the color· pattern.

The indiv.tduals discussed above inwh:toh.:tbe·:r&tioulated dorsal pattern is distinct, usual'.cy- have the pattern arranged so as to sugg(!)st, an outline of paravertebral. light ~s, and often a 'third light line is present on the posterior one thil:'d of the body.· This lattel' stripe is found on the m.1.ddorsal line and oou,:-ses anterior~ £:rem tha eoocy.x • . Speo~ens from Somerset, Texas (KU nos. 20029-42), referable to

.2.• poµol}i on the basis or locality11 lack the reticulated color pattern.,

Instead, ?1Um$roUS small. blaok warts a.re evident on the ·d.orsum. They thereby :resemble certain individuals :fromweatern·¥ted States and western Mexico. However, individuals from Rio Grande City, Texas do agree well w1 th the eubspeo:t:t'io diagnosis ot £• qough:b in possessing a tinely reticulated dorsal pattern, whioh vague~ outline$ a pair of sinuous paravertebral light i:u.a. Reco¢t1on of geographl.Q.ra~es abou1d properly ~ta detailed analytic- atud;y o:r more :material £rom more looalitie:, than. is a:v:~b1e , to me.. In addit:ion, £1$1.d not~s on color pattern will undoubte~ ' \ . ,_. :.-,,.-.,:·.- ,. ,··' . proV$ usef'Ul in the study o£ the speoies; many iulpo3."tant details of oolorat:ton are obscu:red o;- ren<:k,red useless by pre.serva.tives. Zweifel and Norris {lg£•. s._t,.) $'tate. :that their materi.a1 f'x"om three localities - . . . ' .. . :1.n Sonora exhibit <:ertain difterenoes.:1.npa.ttern, 0 but these oannot

· be aclequatel;y demollStrated. wi. tbout oompar:tng living ma.teriaJ..n G@neraJ. gistl:ibut;tgni Sgappi.opus cough\, ocours in southern Arizona and New Mexico• southwests~ Oklahoma• most. of Texas exoept · th$ pa.nha.ndle and extreme eastern part,. sou.thward into sonora,

Chiht\ahua• Coahuila, ?.fuevo Le9n1 Tama.u:J4pa.s., Sinaloa,< ttorthiml Nqant. ' ' . . ', . . . . ' . . Du:rango• Zacateeas, San .Luis Potosi,; and nortbem Veracruz. :rt 1si also found nearl,y' tho entire length of the peninsula of Baja Cali.torn.ta .. .a,. 4ou2}.µ..a according to Bragg (19Wfa.t524), is con.tined to quit$·. xerio ha.bitats. in shol:'t-grass pl.a:i.n$ a.nd deserts. However, along the

wast coast of Mexi.oo this species ooc;urs abu.ndant)Jr in the more mestc Thorn Forest and. T;-opioal Deciduous Fo:r.-eat.

pj.Jzoµssioni Schmidt (19.5:3=.,8) gives the i-ange of'. S~2J?'.M gouol}i as "The Great Pla:i.ns• North Dakota west to Idaho; south to

southern Texas and NW' Mexi.oo." To my knowledge• no valid l:'8QOrda of this species exist north of tat. :,6°N. Records in the literature

of .§• couoh;\ £rom. Utah by YarJ:>ow (187.5:.526j 188:31177~ ,526) and Ga.man 112 98

112

Fig. 2. Geographic distribution of Sca12hiopus couchi. 48.

(1884146) are unquestionably :llt error and properl;v' appl.y to a-nter:nontana. (aee Tanner, l931:171). Attention· i$ dr,m,m to the :tn-. appropriate common name• 0 Graa.t Plains $pa.de:too-r;f•, given .tO'JJ' this speeies in reterenQe to its ~ange by Schmidt (lQs.. git.).

It is &Vi.dent ttom the above :notes tha·t; ail'bspeeifi.o allocation of numerol.Uj specimen& befQre me .is impractioal if the oriteria. l)l"oposed by Smith and Sanders. U2e_,. .9il.) are used. Marked di.ttf)rances in color and size a.re present within and between looa.1 populations in the ·eastern as well_ as western parts of the range o.f i• ~sanol:!¾• In the light of these va.r.tations, whioh are seemingly- not limited by geograpey, it is f'elt tba'l;. separation of subspecies of goaosi ia at .present Ullrta.rranted.

In triy opinion, the dU'ferences heretofo~ used to defi.ne the two Gilb,.. species .of this spa.defoot toad appea:r to be too inconaist,ent to be 9f

Oo. (1 KU); i'1at Santa Mountai;ns~ Co. . near, Rita south. ot Tucson, Pima . (2 KU); 1Jm{ Hexiooi 2 N CJ£ Carlsbad (1+2 KU), 2 mi. 11E of Carlsl,)ad• . . . . mi. . . ' 00. Jl;ddy.~co.. (2 KUH. neax-'Florida,. . tuna (94 rro}J ~om4: T:ll]m;u, oo.

(i KU)J Tern somerset, Bexar Co. (jo KU)J Alpine, Bren-rster Co. (2 KU); Btc»msville, Cameron co. (2 KUH no spe~:l.c; looa.llty. C&tleron Oo. (Z Kµ)J San Diego, Du.val Co. (.3 KUh 5 mi. NE ot Ali.ea, Jim Wells co. (1. KUH Waco, McLennan Co. (4 K.U)J Los Olnlos, :3 mi. E of ru.o Grande Oit.y, Starr co. (28. EHT-m1S)1. Wiobi:ta Falls, Wichita Co. (2 KO); no speoi:tie lo¢a.lit-.y other. than just 11Texastt (4 E!l'r•ffi>fS). MEXICO: -~ Q.al.1ioaa: . 2•.5 mi. SW oi" La Paz (7 STAN)J 5 mio NW of La Rivei--a -(15 '.S'rA'N) I San J'osa' del Oabo (6 STAN); Qoehµilai 3 mi. of Arteaga, .5300 ftc. (2 KO)J Las Margaritas, 2800 tt. (1 KU)I Pan-as, 5470 :rt. (2 lCU)i near Saltillo, Rm. 471. (1 EBT-~H 2... .3 mi. E of Tol'l"ebn (1 WlT-lIMS); }l3lrango: 25 mi.

N of Bermejillo (2 EI!'l'.'-RMS)J near Pasaje (6 Jml'•BMS)J lJµevo ~•

Sabinas Hidalgo (4 m)j ~q.3;1 Maza:tlah (12 EID.'.. BMS); .io m1. NNW ot Los Mochis (1 ItUh $ mi. SSE Rosario, 100 ft. (2 KU); Sgnora.1 0.1 mi. S ot Cooorlt (1 STAN)J h$ar G~a.s {3 EHT•HMS)J IJ.ano (1 STAN)! 2.5 mi. s of IJ.ano (1 STAN)$ 6 mi. s of IlmlO (2 STAN); near Norla (1 :tmr-mts); between Santa Ana •and Llano, Nogales-Hermosillo lu.ghway

(4 S'l'AN)J T,ama.,!J3tlsH!i 1 mi. S of Al:tami.ra (2 KU); 7 mi.o NE Ciudad.

Viotoria (2 KtJh Hacienda. La. Olemantina.1 3½ nd.. W o:r Fori6n (12 EflT•HMSh La Pesca (1 KO)J Soto la Marina.• 500 ft. {1 KO'); zagatega1s to mi. N

Villa cl$ Cos (16 EBT-mw).

GENUS SP.EA

Cope, 1866. Jour. Aead. Nat. Soi. Philadelphia, vol .. 61 sar. 2, pt. 1, July 1866, P• 81 (Generic type, ~oappioww 'a9mbil)'gn§;

Cope.. 186.31 b.r original. d.esignation).

P:lirm

ACOOUNrS OF SPECIES AMD SOBSPEOIES

-~- bgmW,f'Pf?n;t (Copej) (pl. I,· .fig.I\IJ pl. V• fig., 2; fig.; .3)

~(~apbtopqs} l?.,omb~ Cope, Proc. A~ad. Na:~. -Soi. Philadelphia, 1.5t . . 1862 (Maroh, 1863), P• SJ. .Type localitfJ - Fort.. Union, on

Miesoui-1. R:bror, I.at. 48°N; on Platte R1vor 11 • 20 m.Ue;s west o£

. . Fort Kearney; Ua.no Estacado Texa:,; Bouleng~-rll Gatal.ogu.e ot the Batraebia $alientta in the Br.1.titJh MwJeum• 2nd ed... 1882t P• 435; a~, Bull. Eiisoot Inst.; vol. 16, 1884, p. _46 (Mi.ssouri• Nebraska,

Texas)J Sinith and Leonard, .Aln.er. Midi. Nat.,. vol. 1.5 1 1934, P• 190

(Oklaholll9.)J Smith, Amer. Mi.dl.. Nat., ·vo-1. 15 1 _no, 4, 1934, P• 427 (distribution in Kansas); Moo:re, Oopeia, no. 4, 19.37, PP• 22.5.226 (halrl.ts in Oklahoma); Tiben, Trans. Kansas Acad. Sci., vol. 40, - no. 4, 1937, P• 401 {additional looslities in Ka.nsatJ)J Tl:~e and. Trowbridge, Amer. Mi.dl. Nat., vol. 71,· 1937, PP• 460-480 (embryology, liff! ht.stoey)1 Stej11eger and Barbour, Check List- of North American Amphi.bians and Reptiles, 4th ed., 19'9, P• 29J Tanner, Great Basin Nat., vol. 1 • no. 1i 1939, PP• 12.;.13, pl. 1, .f'igs. 11 ... 12; pl. ;3 (Ol"itioa.1. taxonmt4q stu<;\'Y')J Ti.hen.and Sprague, Trans. KSllSas Aoad. Sci., vol. !J.2, 1939, p. ,;01 (Kansas); Bragg,

Ain$r. Midl. Nat .. 11 vol. 24, nc>. 2, 1940,, p. 3:32 (eco1ogiaal distribution in 0l$vel.and County" 0klahoma).1 Bragg, Copeia, no.

1, 19411 P• .52 (Oklahoma); Bragg, wasmann Collector, wl111 411 no. 3,

1941, PP• 92-94 (~sar.tption by Smith (19:34) of tadpoles 0£

bombi.frO!lfl is that of AAfill!lOnc.i.;. and description by Gilmore (1924) and Wright {1929) of hammondi is actually that of bombU;:omih 51.

Bragg• Great l3asin Nat.., vol. 2, 110. 3, 19411 · P• 113 (records b:reedillg 1n day ·~ in New W)x1oo); Trowbriclge1 i't"ans •. Aln~• Miqrosoopt soa., vol. 60, no .. 4,, 1941, pp. ,5os....526 (stud;r ct · dev~lopment of emh:ey-o); Bragg, TUrtox ]fews, vol-. 20, no_. 1& 1942, p. 12 (bteeding b$haV10±-)1 Bragg, soienae, vol. 9.5, 1942, P• 194 (behaviol" in Oklahoma}; Bragg, Proo. Oklaho?na Acad. sci., vol. 22,, 1942, P• :16 (o~oma rseordsJ; ~a.gg a,nd Smith, ct.t"eat Basin Nat., ~l. :;, no., 2, ·1942, pp. 3:3 ..35e. 37-45, 47-49 (breeding behavior in Bragg, Turto~ vol. 20, ,11, 1942, . 9klahoma); ' ' Ni:rwa,. ; . no. p. 1,54 (oklab.oma)J Jtudson 11 Mabra.ska. Cons(lrv. Bull~ no. 24, 1942,

PP• 19.20, p1,, 4 (d1str.tbution in Nehraaka)J Moore and. ni.gney, Proc. Oklahoma Acad. -se1 •• vol. 22, 1942, P• 78 (Oklahoma); i'l'mrbridge, Trans. Anler. M:l.qrosoop. soo •• vol. 61, no. 1,. 1942, PP• 66-8,'.3 (Developm.ent,o:t em~Q and 1a.rvae)i Rod.eek, Univ ..

Colorado Mus. Leaflet no •. 2~ 19'-1-3, P• 4 1 !~• 2 (distribution :tn Oolore.do)i Stejneg~ .. and ·Barbo~; Cheek t.1.st of No,:,th .. Am.erioa.u

Amphibians and Reptiles, .. 5th .ed.J 194;3• .P• 37J Bragia A.•,u~~. Nat• i ·• vol. 78• 1944, PP• $1?"!'.533 (d1.s~bntion in oklahomaJ cntieal study and or w, history); Man-, Amer. MidJ.. Nat., vol. 32, no. 2, 1944, p. 419 (Kans~~- Nebraska); Ande:r$on, Bull .. Ohio.ago Aoa.d, Soi •• vol. 7» no. 5, 1945;, P• 272 .(fust record

.from MistJouri)J Bragg, .Am~. Nat.• vol• 79, 194.5, PP•: 52... 72 · (breeding 'behav:t.o:tl'j phyloger.w)i· Bragg,. He:rp¢tologlca,_vol •. '• 19146, pp. 89-97 (aggregation ·with oannibalism. in tadpoles and its evolutionary signi.tio;mce) l B:reulce~ e.nd Smith, Univ. Kansas Publs. Mus. Nat. Hist., vol. 1, no. !S, 1946• P• 103 (Ka.'Ulas)J Dundee and Bragg• :Proo. Oklahoma Aoad., Sci•, vol. 26, 1946, P• .52.

18 (Oklahoma}& Baxter• W;roming Wildlitei. vol. 11~ no~ a, ~947, P• :33 ( distribution in Wyoming) J Bragg• Wa.smann. Collector,, v<>l. 7 ·no. 2, 1947 • p. 71 (aggregational phenontena in tadpo~s) J Bragg, Proo. Oklah~ Acad. Soi.., vol. 27, P• 61 (tadpo1e behavior); Li~ze,- and Wright, •r• Mi.dl. Nat.,irol~ 'J7,, 1947, P• 202

(eggs desox--J.bed) ;. Wheeler, Alne:r. Midl. Nat., vol. ;38, no. 1, 1947 t

P• 1'7.3 (distrl.lmtion in North Dakota); Braggj· Wamnatin Co'l.leotor, vol. ·7, no. 2, 1948, P•' 71 (tadpole b@harlor)l Bragg, Proc. Okla;;. homa. A¢ad. Soi.,. vol. 29• 1948; p. 11 (~ seoretiohs by femai~s

. suggests sexual di:ti'erence); Bragg and Dimd~e, Proo. Okla.hem.a. Aoa:d. sci.; vol. 29, 1948, P• 2,5 {Oklahoma +'$Oords)J Bragg"' Researches on the Autphibia o! Oklahoma,,· 19.50, PP• 26; 3.5, 43, 78 (distribution, breading habits, and adaptations 0£ survival. vaJ.ue

in Oklahoma); Bragg, wasmann Jou:r. Biol.; vol .. ·8, no. 2., 1950, PP•

221-228 ($o

Oklahoma Acad. St;d., vol. 30, 19.!51, P• 20 {Oklahoma records); Stebbins; .AlY!phibians of' 1-:estetn l~orth knerioa, 19.51. pp. 202 ... 207, pls. 14, 271.1 2s • .55 (genera1 diaoussion); Axtell and Wasserman, Herpetologioa., vol. 9,; 195.3, p. 1 (extension cit known i-a:rige into

southern Texas); Soh:midt• Check list of North .American .Amphibians

and Reptiles, 6th ed.,, 19.53, P• 59) Shannon, Herpetologioa.11 vo:i. .. 9, no. J, 1953, PP• 127.,..128 (f'irst def'ini.t.~ reaord.:trom .Arizona.); Tinkha111, Herp&tologioa, vol. 8, no. 4, 19.5.3, P• 179 (prey item of "prair.1$ rattler" !n North Dakota); Bresler and Bragg• Copej.a, no. 4• 19,54, P• 2,56 (variation in rows of labial teeth of tadpoles);

Orton, Oopeia, no. 2, 1954, pp •. 97.,.99 (dimorphism 0£ larval mouthpa:rt$ discussed) l stebbins, Junpbibia.ns and reptil.~s ot

we.stern North America, 1954, PP• 9,, 141• plso 11, 17; fig. 13 (general disCU$s1onh Tihen, Copeia, no. 3, 19.5l}, PP• 217.. 218 (fossil material trc>m PJ.eistoceue of Kansas rei'etred to this

speoies)J Wheeler11 Amphibians artd ~ptUias of the North JJakota Badlands, 19.54, pp. 14-16, figs. )-4, 11 ... 12 (dist:rl.butie>n and general account); Blair, Texa.$ Jour. Soi., vol. 7, no. 2, 1955, P• 186• :r1g. 1 (description of call).J Du.ell.man, Occas. Papers Mus. Zool. trr.d.v. Ml.ebigan, no. 569. 195.5-. P• 2 (Arizona); Logior and Tone1"JI Oontrlb. Royal. Onta.:rio Mus. Zool. Pa.leo., no. 41, 195.5, pp. 22-23 (dist.rib ution 1n Canada); Smith. C:opeia, no. 2, 195.5, P• 1)5 (reports on a pallid speo:bnen from Ne-.r Mexteo)J Walt&rs. Oopeia, no. 3, 19.55, W• 2.52-2.53 (A:rrl.zona); Zweil'el,

Amer, Mus. Novltates no. 1762 11 1956, PP• )S--36, figs. 21 ... 2.3

(pl\vlogeey and distribution)., s:e2a bombi:fl;:oM Oopa, JOllJ.-. ,A.cad. Nate Soi. Philadelphia, vol. 6, eer. 21) 1866, P• 81; Allen, Proo. Boston Soo .. Nat. Hist., vol. 17,, 1874, P• 86 (Wyoming)J Cope, Bull. U. s. Nat. Mus., no. 1• 187.5,, Pe 31 (11$1:.ed); Amere Nat., vol. 1:3, 1879, P• 436 (Idaho, Montana); Yarrw, Bull. U~ s. Nat. Mus •• no. 24. 1883, P• 177 (listed); Brown, .An annotated eheQk list o:t the Reptiles and. Amphibians of

Texas, 19.50, P• 40 ( distribution 1n Texas)& Firaahein1 Herpetol• ogioa, vol. 6, 19.50, pp .. 75•76 (first reQord .front Mexico; state of Chihuahua) s Smith, Handbook of amphibians and reptiles of Kansas 19.50,·pp. 66-68 (distribution 1nKa.nsaa; general ac11,1ount)l Bragg and Hudson, Great Basin Na.to,, vo1. 11, nos. 19.51, P• 87 (Oklahoma reoords). 54.

lgynmgndi ]2omJ;nttons Cope, Bull. u. s. Nat. Mus., no. 34, 1889, P•

,306, fig. 78,·pl~ 51 1 fig. 17& pi. 68 11 fig. 1a; Cope, Proo. Acad. Nat. Sci. ·phUad$J.phia, 189Z, P• 333 (Texash· Co~, Anier .. Nat.•

vol. ;30 1 1896, P• iOi6 (geographical distrtbu.tion}J Bailey. North

Amer. Fauna, no. 3S, 1913, P• :35 ("Upper Sono~a.n Zone in New . Mexico")• .S,op.~s hglondi,: Gunther, Biologia Oen~ali-Amerioa.na'.. Reptilia and Batl"achia; 1901,' P• 2.59 ~-h Ellis' and Henderson, Univ.

CoJ.ora.do Studi.Eu.-;, vol. 101 no.- 2, 1913, PP• 5:t.-.52 (distribution in Colorado); Stejnegel." and. Barbour, Check t1$t of .North Amerioan

.Amphibians and Reptiles. 1st ed., 1917, P• 25 ~-)J Nieden, l)as. Tierreioh, Lief. 46J Anu.ra I, i923, p. 47 (;gart.); Stojneger and Barbour, olieck'l.1.st ot North.A:mer.tcan .Amphibians and Reptiles, 2nd · ed., 1923,>P:~ 2) (;Qart.)l Young, Quart. Jou:r. Univ. North Da.kota1,1

: vol., 15 11 1924, P• 59 (North Dakota); Storer, Univ. Cali:torn;i.a . Publs. Zool •• vol. 27, ·192.5; p. 148 (general discussion) ~.)J

01-tenburger, Proc. Okl.ahoma Aaad .. S

(Oklahoma); Oookerel.l,. Zoology o:t: Colora.do/1927, P• 112 (Colorado)J Slevin~ occas. :Papers Qali:f.'orxxia Aoacl. Soi., no. 16, 1928, p. 84 (ganara1 discussion) Cll!r.t.•H wright• I>roo~ u. s. Nat. Mus,., vol.

7411 1929, PP• 4, 18, pls. 11 6 (tadpoles of' bgJnb1Jrops described and figured under name of 1¥Wimondi)j stejneger and Barbour, Chec,k List of North .American Amphibians and Reptiles, 3rd ed.1,1 19:3:3, p •. 26 ~-H Wright and Wright, Handbook of )Frogs and Toads, 1933• PP• 40-41 (md.) SoaplJiopus llm9mongi gbitrnps, Dickerson, The FJ!"Og Book• 1906, Pe 61, pl. 9 (habtts, des':::riptioli, distribution); stre1:;ker, ~c,c,.

Bi<>l• So<'-. Washington, 'VOl. 21• 1908, Pe 60 (

vol. 13, 1910 1 p .. 17, .fig._· 1 (no:rthwestern T~s);_ Dickerson,

The Frog Book, 1913, P• 61 1 pl. 9 (genaral dtscusaion)J Cary, . ' . North kner:lcan Fauna,. no. 42, 1917, P• 19 (1ooalitie41 in Wyom1t1g)J Over, Buil. SOtlth Dakota Oeol .. t-Jat. Hist. SUrv., no .. 12, 1923~ p., 12. pl. 1 (distributiuth Dakotah Gilmore, Colorado

Coll. Publ. _. SOia ser., voJ.. 13,no.1 11 1924, PP• 1-121 figs • . ' . . ' . ' . 1-7 (lif'e history st'Ud,yj disttlbu.t:i.on· Colorado); Burnett, •oooas. Papers Oolora4,o state Agr. Oofi. ·Mua. Zool. Ent., vol •. 1, nc>. 11 1926,. P• 2 (Co1ora.do)j Tayioi6, Uni~e- Kansas Sci. Bull.,

voi. 19t no. 61 .1929• P•. 6,5 (~~st tieaord .from Kansas)j R:ell.ogg,; •;Oo;peia• tlQ.1,.19:32• P•. 36 (l.toniaha)t Br$nnan• Trane. lfansais .... · . Adtidi._ Soi.., vol. :37, 1934,· P• 189 (Kal1sas)a Wright and Wright, ( . . . ·. -~ . ' .

. . . ' . . . . ' . . Wrl.ght• ·• (general disoussion)f ·.· · 1949, ~P• 116-±20,. ;pl~·: 19' , ' •,) ' . '· Moore• Oopeia• .no. 4, 19.52. P• ·27fl (distribution .Albet,tai ·· · ·. Canada); Mosimann and Rabb; Oop¢~a. no. i, 19.52, P• 24 (Montawi)~

·?Sqa:rab3:opw1, Deokar• Proo. Okla,homa Aoad •. Soii-,.. vol. 10 11 19:30; pp •.·. ;. . . 82-84 (spade£oot. 5.mpri.aonad tti ba'l.:lJj of ~la.;r below surface.· in· . • . ,r '·· Oklahoma); Stabler• .Ht,rpeto:togica• vol. 4, '1948, P• 168 (prey itEml of Crotal}l§ in easuji!n• Oi;>lOl:'ado)~ . . ' Taxgnomiq W:aio,r.y;a Cope (186);.SJ) d~sm.beithis spe.cies fl-om specimens in the U. s. National Museum taken by l'il"• E. J • Pennis at ''Fort . Union• on Missouri River• 1at. 48°N.u (no. 1704); 'by w. s. Wood. "-on Platte River, 200 mUee,west.of Fort Kearney" (no. 3.520)i atid by;? capt. Popsits 'Expedition trom JOI,la.no Estaoado Texas'' (no • .3703), Schmidt (19.53:59) restricted the type 1ooality to Fort Union (wUliam.s Co.), North Dakota. Thia spadef'oot was designated the.· ·type species ·of the •·aenus by Cope (1866181). In the same pa.PE¢ ru,. assigned two oth&:r :speaies to the new genus. Later Cope {188Sh)04) noted intergrading· trends between Jlti.Jm. hammonsii and ,a. pom.bifroa, an(l ~uoed both to $U'bspeoitie 'rank. Since then the status of pompi.frors bas lortg ~n o~ o:t diaput~. S«iveral • . . : •' . . authorities (Dickerson, 1906, 1920; Stejm,ge:r:- and Barbou.t.;, 1917, 192:3; . 19331 Goldsmith, 1926, Slevin~ .1928; orien\rorger and. Free~, 1930-,

' ' Kellogg; 1932a; and others) have ~<>nsi~red it a synonym o:t ii• h~opg%.

' '' Sm1.th (19;34s4:34) was th~. :ti.rst to qlearly dernon.,trate the speoitio · inclependenoe 0£ bombi££2m• Tanner (19.39). in his criticai study of Bragg the Amerioan spad&f'oot toads, treats it aij' ...4• full speoies. ( i94ib) found no di:t.ticulty in dietingu.tahing the two specie$, lW;mmoAd\.. and.l?9mb1£rons, on tho basis oi' oalls -a;;:_.woll as the st.X'Uc:ture ~t·larva.e and adu1tsti With few exceptions (i~ ••• wright. ·and Wrl.ght. 1942~ 1949;. Moo~, 19.521 Mosimann and Rabb, 1952) all su.b$ag_ti13nt ~rket'S have tollowed. . Smith and Ta:tU1er. treating thia spade£o6t .as a distinot. species,. _

Diagno§i;; A~ with a_·fron·toparietai·tonta.ne~~ a di$tinot bony intororbital boss; dorsum relatively smooth• ,dth tw small red-tipped (in li£e) tuberalesa dorsal color pat.tern composed of irregularly arranged darkblotohes of variable shapa on a dark grtq background; tympanum small• often indistinct; parotoid usually indistinct s spade slightly longer than, or as 1ong as wldel. ya.riatign: No consistent variational trends were noted. However• three specimens t'rom Navajo doun-cy, .Arizona (FAS nos. ,'3013; 3577, 3.579). differ :from typical spe

1.)lontans ·may b$ a subspecies of ~• ggrnbifrolll.• He was probably influenced by the above three speoimens ~rhioh :resemble g. intermont&J.1! in possessing a large parotoid and tympanum and numerous large dorsal tubercles e But on th& basis ot the bony interorbital boss I :refer them to §.~ bombi.tronq.

The three individual.ti mentioned above constitute an extension 0£ the known t-ange northward nearly :,oo miles from Willcox, Coobis~ County, Arizona, the northernmost looa.11.ty 1n Arizona reported by Shannon (mi.. c~t.}. He was the first to report this spadefoot .trom Arizona. Distripu.tiona l?9mp1:t):0M, ranges .trom southern Alberta and extreme southwestern Saskatchewan, Canada southward into Montana, North

Dakota, eastern Wyoming• South. Dakota• Nebraska, Kansas. eastern Colorado.,

Oklahoma., the Texas Panhandle, New Mexico, southern and extreme northeastern Ar.bona. and northern· and northeastern Ohihuahua., Me:x:1.co. Axtell and Wasserman .(19.53i1) reoently reported the presence of this spade.toot in extreme southern Texas. Alt.hough records are still lacking tor Mexican st.a.tea othor than Chihuahua., it seems reasonable to assume that the range of _a. bombil'ronm is oontiriuous across northern Ooahuila. and Nue\"O Lebn to southern Texas. 34

\\5 58.

P?-soussiont w. Frank Blair in.forms me (in~.} that he bas . . ' found hybrids between bgmbi.frons and· Q@llOQ at two loqalltiea. one in Trans-Pecos Texas and one in the Texaa Panhandle. However, he regards these two· as·. good epeoies wbioh .show limited bybridization11

This speciel9 was firet recorded £ram Mexico (st;atE) of Chihuahua) by Firschein (19.50:?S). In the Univeraity of Kansas Museum of Natural History there are thirteen specimens or ·bombigrons from two looalities in northeastern Obihuehuawhioh provide the third and fourth

record stations 0£ occurrenoe ·in the state and oonsti.tute an extension·

' . o:t the known range for the speoies nearly 150 mile$ southeastward in ... Mexico from the nearest previously reported locality• ·Chihuahua, sta~

ot Chihuahua;. reported by Stebbins (1951:202).

Ti.hen (19541217) refers one ilium and one fragmentary sao~al vertebra with the -anterior portion ot the· tusad coccyx from the Pleistocene of Kansas to this species .. According to Bragg (1941b:92) the tadpoles ot J?omb;t.rrons ware deaorlbed by Wright (1929:4_ 18, pls. I, VI) under the name ham,mondi. ·

The description 0£ tadpoles ot bombib:gns by Smith (19.)414291 pl. XII) . properzy applies to llammopdi• Bragg Cloe. ~.} adds• Speoimem examinegz 765; ~i Red Lake (2 FAS)- Shonto Canyon,·Navajo Co. (1 FAS); 13 mi. E o:t E:U-rida, Coo~se Co. (2 FAS)I Kansa.sa 1 mi. Si of Aetna (23 KU). J/4 mi. W 0£ Aetna, Bar~r Co. (1 KU); Stephenson Ranch, Clark Co. (2 KU); North Lawrence (22 KUH

3 mi• N and 1 mi, E of tawrenoe1 Douglas Co, (1 KO)J -Garden City; Finney Co. (7 KU)J i mi. E 0£ Coolidge, Hamilton Oo. (1 Kt7); Harper,

Harper Co. (1 KU)J Rezeau Ranoh, Kiowa Oo. (1 KU)J 1 mi, Wot State Lake (2 KO), Meade County state Park, Meam, Co. (1 KU); 18 mio N of El.kart

(99 tro). 18 mi. NE o£ El.kart (1 KtJ) 11 no spacifio locality, Morton Oo. (1 KU); Medora (:; KU), near sterling (1 KU). ho apeoi.fig looality• Reno Co. (2 KU) J Nekoma, Rush Oo. (134 KU)J 4 mi. S of Scott City•

Scott Co. (2 KU)J i mi. E of Liberal (3 KU}, 4 lllio N o:t Libaral, Seward Co• ( 1.5 KUh Goodland (1 KU), 1 mi.. S of Goodl.and {47 lCO'), no specif'io locality, Sherman Co. (1 KO')J 11 m1. E o'Z Hugoton, Stevens

Oo. (4 KO); .5 mi. S ot Brewster. Thomas Co. (1.3 KU);· Laoey Ranch,

Wallace co. (1 ICU)a Nebr3akai 4 mi. s 0£ Olay canter. ·OJ.ay- co. (1 KU) J 14 mi. 8 0£ North Platte, :tinooln co. (1 KO)J 4iew Mextco;

Melrose (64 KU) 11 Claude (36. KU). Clov.l.~, Ourry Co. (182 KU)J 10 mi.

SE o.:r Silver City, Grant Oo. (1 KU)J near Clayton1 Union Co. (1 KO)J near Inscription Rock (2 KUh near Grant, Valenoia Co. (1 KU)l

Oklahoma& 7 mi. S of Boise City, Clil.n.~n Co. (1 KU)J S mi. W of Cantons; De-111ey Oo. (17 KU); along highwq 10 between Braggs and.

Muskogee, Muskogee oo. (1 KU); Clevela."ld, Pawnee Co. (6 KU); south Dakota,: Buffalo, Harding Co. (.5 KU)J Verntillion, Univ. South Dakota campus, Oley Co. {8 JSF)J bxaa: ? tni.. S>l 0£ Byers, Cla.y Co.(1 KU); 5 llli. E c,£ Cacy-on, Randall Co. (7 KU); WX

ha.nnnon9is (Baird) (see rsynorrpi:r under subspecies) l'l§U¥!0~ ranges frOlll westarn California southward into northern Baja Callfonda, eastwa.i'>d. through Arizona, New- Mexteo into extreme western Tex.as. uorthw.u-d 'into nortbweste:t".n Oklahoma, western Colorado, an4 northwestern Wyortd.ng. Tanner (194-1*88) reports the presence of this species in northeruJtern Washington and .northwestern Montana., but gives no definite looalities. In Mexico 2.i. h!mmon91 is found in nearly all the states. ~eacbing its sou.them limits' in·central Oaxaca. Exand.nation of material. from throughout the entire range.or·~•

hamm9ndi ::reveal.a the ~Xistenoe of $. de:tinite north-south allne. ·The Oline is evident µi size• the·varying·(legree$ of.dorsal i-ugos1ty, the

ratio of width to length o£ the.spade, the length o:t th9 hind leg

relative to the total length of the body• and the degree 0£ develop..

titsnt 0£ the pa:rotoid gland. The sou.the~ subspecies, !!m+'tiplioatfb is g(!neraJ.l1' a smaller

animal (max. snout-veilt length reoorded .59.0 mma J av. 47 .,8 mm.) than the northel"Il form (max. snout-vent length :re,;,orded is 62.4 mm.j av. 49.6 mm.). has aoa.rsely tubel:'oular dorsal skin (relative:cy- sxnooth in h• bammon~) • a spade usual.lY' as long or longex- than wide (generally wider than long in the northei-n subspGa:Les);. and a well-developed.; though small• parotoid. gland ( this gland 1s usually indistinct in h• hamm2a<~J. In addition multi:QUcato!\ has a slightly longer hind leg

than the northern subspeoies I th& spade rea.obing to between the tympanum and anterior margin of the eye (when the bind.leg :ts brought forward . parallel to the body). In the northern :f'orm the, t1pa.de scarcely reaches 114 98

• specimens examined 22 o other specimens

0 400 I I I I I scale of miles

114 98

Fig. 4 .. Geographic distribution o:f Spea hammondi.

1 • s2ea h. hammondi 2. Spea h• multiplicata 61.

the tympanum. It. is possible that with additional material from critical areas

/ ' ' / in Queretaro. San Luis Potosi• and Verao:ruz, other geographic races~ be separated. The variation of individuals from these areas 1s discussed further under hamptoncMi my1tj.p)4cata.

AAID!BO~ ham!llondi (Baird) (pl. I,. fig. 51 pl. IV, :f'igs. 4-.5; £ig. #) ScauM,opµs hamon4\: Baird, Report._ of the Pacific Rail Road survey, vol. 10, Will1amson•s route, pt. 4, no. 4, 1857 (18.59). p. 12; pl. 28,

figs. 2a-2d. Type locality: Fort Rea.ding, CaliforniaJ Cope, Proo. Acad. Nat. Sci. Philadelphia, 1863, P• 53 (description); Cooper, in Oronise, Natural Weal.th of California, 1868, p. 486;

Cooper, .Proc. California Aca.d. s01.,. vol. 4, 1870 (1868-1872) 0 P• 64g Boulenger, Oata1ogue of' the Bati-achia. salientia 1n the British Museum, 2nd ed.• 1882, P• 43.5; Garman, Bull. Essex Inst., vol. 16, 1884, p. 46 (listed); Cope, Bull. u. s. Nat. Mus., no •

.32, 1887, P• 12 (~ stagnal;\.;i and §oapWzopus dyge§ii placed

into the synonymy of Scaphiopup halnnton4\)J Stejneger, North

American Fauna, no. 7, 189.31 P• 222 (Cali£orn1a); Gunther,

Biologia Centrali-Americana, Reptilia and Ba.tracbia; 1901, P• 259 (gives range as •'Missouri to Mexico") (~.); Dickerson, The Frog Book, 1906, P• 59 (genera1 discussion) (~.)J Strecker• Proc. Biol. Soo. Washington, vol. 21, 1908,- P• 6o (Texash Cary,

North American Fauna, no. :3.'.3, 19111 P• 27 (Colorado); Bailey, North .American Fauna., no. 35,. 191:3, P• 35 ("Upper Sonoran Zone in 62.

New Me:Jdco")I streoker, Baylor Univ. Bull •• vo1. 17, 1915, P• 54 (two new looallties in Texaa)J Stejneger and Barbour; Cheok List of North American Amphibians and Reptiles, 1917., p. 25J Schmidt, • Bull. Arn.er. Mus. Nat. Biet., vol. 46, 1922, P• 6.33 (first reoord from Baja California.); Nieden» Das 'l'ierreioh, Lief. 461 .Anura I, ' ' • I 1923, P• 47 ~.); stejneger and Barbour• Check List of' North American Amphibians and Reptiles, 2nd ed., ·.1923, p. 23; Cowles, Jour. Ent. Zool., vo;. 16, 1924, P• 107, 5 f~s. (voice, breed- , ' . ing habits .in Arl.zona; tadpol.e £~}~. Ort.en.burger, .l'.roe. Oklahoma Acad. Soi .. , vol. 4, 1924, P• 19, pl... 1 (~bit~ in . ,• 'I •· < . ' ' ' Arizona; eggs desetribed) I Van Denburgh,. Prc>c • ,California Acad., Soi .. , ser. 4, vol. 13,. .1924, p. i_94 (distribution in New Mextco); Englehardt., Copaia, no. 149, 1925, pp._91-92.(b~eding. habits in Arizona)J storer, Uni:v. Cali:f.'ornia Publs. zool .. , vol .. 27, 192.5,. PP• :;6; :39, 148, pl. 10. ti.gs. w. EE (general. discussion; eggs and tadpoles figured) I Goldsmith, Carnegie !z:ist. W~sbington Yearbook, vol. 25, 1926 (192.5-1926), PP• _36~-,zo (h!l.bits and reaatio~a Ortenb~r and Ortenburger, :E>rooo Oklahoma Ace.d ..

Sci., vol. 6, 1926, p. 102 (habits in Arlzona)J Barber and King0

Publ. Health Rept. 11 vol. 42, no. 52, 1927, P• 3189 (larvae feeding P• (Okl.a- f 46 no. -, . O:rtenburger, Copeia, 163,.1927,' on mosquitoes)t' . homa); Sltv1n, Occas. l>.apers California Acad. 501., no. 16, 1928, P• 84 (general discussion;. distrl..bution. in western United states}

pls. 1, 6 (larvae described and. figured are those of• )3orneM't9rm.} I

Myers, Proo. Biol. Soc. Washington1 vol. 43, 19;0, P• 61 (California. records); Noble, The Biology of the Amphibia, 1931,

PP• 403, 406, 408, 415, 421 (general discussion) J l3rues, Proco

Ame1•• Aoad. Arts so1 •• vol. 67, no. 7, 1932; p. 28~ (larva& from hot springs in Wyoming); King, eopeia, no. ~b 19:32, P• , 17; (Arizona)I 10.aube):i, Copeia., no. :3, 1932, P• 119 (Calii'• ornia) j Lins~, Univ. Calil'orma Publs.· Zool., vol. 38, no. 6, 1932, P• 350 (Baja Calitorm.9.h 'Bo.rt, Amer. Ml.dl. Nat., vol. 14, no. 4, 19,33,P• j,50 (Arizona,' Calil'ornia~ Colorado);

Campbell, Copeia, no. 2 11 1933, p. 100 (Arlzona)J Stejneger and

Barbour, Check List ot North American .Amphibians and Reptiles, lrd ed., 1933, P• 26; Wright and Wright, Handbook of Frogs and Toads, 193:3, PP• 40-41, pl. 7 (~.); Klau.ber,. Bu.11. Zoo1. Soo. San Diego, no. 11, 19.)4, p.' $ (Calltornia); McKee and · Bogert, Copoia, no. 4, 1934, p;i 178 (.A..rJ.zona)'J Smith, Amer. Yddl. Nat., vol. 1.5t no.· 4, 19:,4, P• 434 (bammondt distinct from bomb;Y;rgpsi larvae desoribed under name oi' .bombtt')yps);

Wood, Copeia.1 no. 2, '19.35 1. pp. 100-102 (Baja California; Utah,, Arizona) (parl.); Gloyd, Bull.. Chicago Aead. Soi•• vol. 5; no. s. 1937, :P• 103 (Ari.zona)f Little and Keller, Copeia, nc. 4, 1937, PP• 218•219 (Nett Mext.co)i Wl"'lght and Wright, Trans. Texas

Aoa.d. se1 •• vol. 21, 1938, P• 17 (dist~bution in Texas); Smith, Ann. Carnegie Mu:., vol. 27, 19:39, Po 311 (Baja Oallf'orniah

Stejneger and Barbour, Checm Li.st Qf North American Am:phibiazw and Reptilas, 4th ed., 19:39, p. 29 (part.)& Tanner, Great Ba.sin Nat., vol. 1, no. 1, 1939, pp. 16-18,, pl. t figs. 7.8; pl. 2, fig. 1J pl. 3 (critioa1 taxonomic study}J Bragg, wasmann Collector, vol. 4, no. 3, 1941, PP• 92-94 ("Smith'$ (1934) description of 64.

tadpoles qf b_ombif'ropq is _that. of hmmpop.ga.. and descrl.ption by

Gilmore (1924) _and Wright (1929) 0£ hawnonst,. is aot~ that 0£

b011lbi.frgn§"}s Bragg, Great Ba.sin Nat., _vol. 2t no. 3, 1941. P•

112 (notes on tadpoles £ram New Me.."d.ao); Bragg, Turtox Newa 1

vol. 20 1 no. 1. 1942, p. 12 (breeding behavi~r)~ Bragg and Smith 11 Great Basin Nat.• vol .. ·3, no. 2, 1942, P• 37 (breeding behavior i.n Oklahoma); Bragg, 'l'urto:x: News, vol. 20; no. 11, 1942, P• 154, . . (Oklahoma)J Wright and ~ht• Handbook of Frogs and Toads, 1942, I • ' , I PP• 48-49, pl. 7 (general 41.acussion)I tcau£f'eld, Amer •. MicU. Nat., vol. 29, no. 2, 194:3, P• :;42 (~ona); Rodeok• Univ. Colorado · Mus. Leatlet no.·. 2, 194:3, P• 4 (hypothetical list ot Colorado.); ' . . _· ' . . ; Stejneger and BarboUl.", Check l,ist ot North Amer:toa.nrJunphibians and Reptiles, ,5th ed., 1943, P(37 I Bragg,. Amer. Nat., vol. 78,

-19441 PP• 517~.533 (distrl.bu,tion in Oklahom.aJ Qri:M.cal st~. and s'tllml!ary of knowledge of ha.bi.ts); ~rs, Copeia, _no. ,:t. :i.944, p. sa (distribution in California); Bragg, .Amer. Nat.; vol. 79,

1945 1 PP• .52.72 (breeding pa~ternsj peylogen;y-)1 Fitch and Ttd.n• ing* Copeia, no. 2p 19~, P·• ?O (P~Y'. item of QEota.l~s mk1a oreganusH Baxter, Wyoming wUdlite.

behavior); Piaknellt Amphibians mici Reptiles of the F~oifio Statest , 1947, PP• 14• 168, 174, 178, pl. 67 (general disoussion); Bragg.: wasmann Colleotor, vol. ?t no. 2, 1948, p •. 71 (tadpole behavior); Smith and Taylor, Bull. u. s. Na.t. Mus., no. 194, 1948, P• ;36," (distribution in Mexioo)J Jameson and Flury, Texas Jour. Sci.,

vol. 1, no. 2, 1949, P• 60 (Texas); Bragg11 .is Reaearohes on the

Amphibia of Oklahoma, 1950 1 PP• 27, 43, 78 (distribution and

breeding habits in Oklahoma}; Burgess, Oopeia.• !lO. 1, 1950, PP• 49-.51 {development of larvae under laboratory oond.1.tions); Lewis,

Herpetologioa. 11 wl .. 6, 19.50, p. S (reports white tadpQle £rom New Me:d.co)a Stebbins, Amphibians of western North kneri0a, 19.51, PP• 207... 217, pls. 14, 27, 28, .55 (general. disoussi.on) (~.); Blair, Texas Jour. Soi .. , vol. 7, no. 2, 195.5, P• 186, fig., 1 (description and sonogram o:t call); Du.ellma.n, oocas. Papers Mus. Zool. Univ. Michigan, no • .569, 19.5.5, P• 2 (A'rizona); Walters, Oopeia, no. 3, 19.55, PPe 252-253 (Arizona.); Zweifel, .Amer. Mus. Novita.tes1,1 no. 176211 19.56, PP• 35, 42, figs. 11-19• 21t 2J-24 (general dieou.ssion; phylogeny). [email protected]• Cope, Jour. Acad. Nat. Sci. Pbiladelphia.• vol. 6, ser. 2, 1866, PP• 81-823 Cope, Proc. Aoad, Na.t, Sci. ~Philadelphia, vol. 18,

1867 (1866)'~ P• 301 (specimens from "Territory ot Arizona0 )J Cope, Bull. U, S. Nat, 1-{Us.,. no. 1t 187.5, P• .'.311 Couest $Un', W100th Merid., S, 187.S, P• 6:,o (NEM Mexico)J Copa• Proc. Amsx-. Philosoph., Soc., vo141 18, 1879, P• 264 (Guanajua.to and Chihuahua, Menoo)j Cope, Proc. Aca.d, Nat. Soi., Philadelphia, 188)·, PP• 14, 32 (Net-r Mexi.co, Oall.tornia); Yarrow, Bull, u. s. Nat. Mus.it no. 24, 188.'.3, P• 177 (listed); stejneger, North .American Fauna, no .. 3, 1890, P• 117 (Arizona.); Stone, Proo. Aoad. Nat, Soi, Philadelphia, 1911,- P• 66.

22j (N$--.i Menco) 1 Sl'lii th, Handbook 0£ atnP,hibi~ and ~eptiles of Kansas, 19:;o, P• 307 (cypothetieal list of Kansas).

n;taigpalis Cope, Ya..r.row, SUrv. W 100th Mer.ld., .5• 187.5 1 P• 52.5»

pl• 2.5, .figs. 6-8. Type· localityi Alto dos Utas,. northwest Ne-vr

Mex:1.cot Ya:rrow- and Renshaw, Ann. Rept .. Obie!" 0£ Engineers :tor ' : . ' ' ! •

1878, SU1-v'. W 100th Merl.d., App. NN, 1878, P•, 209 (~ornia.) i Yarrow, 'Bull. u. $. Nat. Mus •• no. 24• 1883, P• 177 (listed); Garman, Bull• Essex l:nst., vo1,. 16, 1884, p. l.16,. (listed). §gauhioW§ stagnalif• Boulenger, catalogue·~ the Ba.trachia Sallentia

in the British Museum, 2nd ed., 1882, P• 4:;6. h\\UIP'.!!2.llst\ htgmnondi. Cope, Bull. u. s. Nat. _Mus._, no. 34, 1889, P• 30.5, fig. 771 pl. 46, fig. 8J plo 49, :fig. 18; pl. 68, :f'ig. 11

Cope 11 Amer. Nat., vole 30, 18961 Pe 1019. (geogl"aphical. distribu-

tion); Brown; An annotated obeok 11st. 0£ the reptiles and amphibians of T~s, 1950, P• 4f {distribution in Texas)• .

SoaphioJi>wi b@.!ntnondi nammoncli, Grinnell and Camp1 Univ. California

Publs. Zool., vol. ~7, no~- 10, 19171 P• 140 (distribution in.

Oa.li:f.'orn1.a); Smith and Leoruu-d 11 ~er. M:1.dl. .. m~t., vol. 1.5,

19341 P• 191 (Okhhomah Chao Liu 11 Proo. Boston Soc. Nat. Hist .. • vol. 41; no. :3, 1935, P• :30 (vocal sac described); Livezey and. Wright, .Amer. Midle Nat •• vol. 37; 1947, PP•: 192, 202• .:fig. 5. (eggs desol'ibed and :f'igured)J Tanner, Great Basin Nat., vol. 9,

nos. :3-41 1949, P• 47 (.Arizona); W:right and Wright1 Band.book of

Frogs and Toads, 1949 11 PP• .11:3 ... 116, pl. 18 (general diE>cussion);

Turner, Copeia• no. 3, 1952; l'P• 172-175, 2 figs. (~scusses variation in inouthparts ot tadpoles)J Obilds,Evolution, vol. 7,

no. :;, 19!5,, pp.• i-;228~233 (predation by raoooon on albino tadpoles

. ' in Ca.li:tornia)j Sohmidt, Cheok_List of North A,merlcan .Amphibians and ReptilGs, 19.53, _::,ii.h ad., p .. .59; Ovton, Copeia, noo 2, 19.54,

' ' PP• 97•99 (dimorpbi.sm'in larval. moutbpa.rts diacussed)J Stebbins,

Amphibians and :reptiles o! western North Amerioa1 19,541 PP• 93 1 143, pl.~~17, fig. 13 (general a.oaount.J aistrtbution); Murray, Herpetologioa, vol. 11, no. 1, 19.55, P• _j3 ·(Baja Cali£ornia). a,caph12PJ1S )?ombi.f't2Jli, Stnith~ Alner~ Midl., No.t •• vol. 1.5, no. 4, 1934• P• 429, pl. 12, fig. 1 (mou.tbpal"'lis of tadpole of hanp:nondi described and ~iguredunder nawt o£l?s?mbitrons). h.a.mmondi mµJ,t;l:.pligata, tangebartel and Sim.th,- Harpetologioa, vol.

10, no. 2, 19.54, P• 125 (second known locality in Sonora) J

Ohrapliwy and Fugler11 Herpetologioa, vol. 11, no. 2, 195.5, P• 121 (two localities in Chihuahua, Mexi.oo). '.fgopom.tq hi;tottt Baird (1859:12) described aoapbiopua !19on,

Diagno§~§t A Im!! with £aw small scattered dorsal tuberalesJ spade usually wider than longJ pa.rotoid gland generally indistinct; tympanum. usual.l;r indistinctJ spade soaroely reaching tympanum; no bony interorbital boss. 68.

J[aria.tiont Individuals £ram throughout Calii'ornia are large (max. snout-vent length recorded ia 62.4 mm.; av• .53.:3 mm.)-, have a relative4'" smooth skin and possess a large tympanum (it va.t'ies .from 4.0 mm. to 4.6 mm. :tn diameter) the diameter of which is usually greater than tha distance :tram the nostril to the eye. The parotoid is indistinct and the spade is blunt and short. being mu.ch wider at its base than long. The internaria1 distance varies frQJ:?l :3.1 mm,. to 4.2 m.m. (av. J.6 mm.) and the interorbital distance varies from ;3.6 •• to 5,.5 mm. (av. l}.7 mm.). The spade scarcely reaches tha tympanmn when the bind leg is brought forward parallel to the body,. One exception to the above is an individual f:iom Sierra Madre, Los Angeles Co. (MVZ no. 4,3;4). It has numerous large dorsa1 tubercles, th$se more concentrated dorsolaterall,y.

A large parotoid gland partial.4r covers th$ posterior margin ot the small tympanum.. In all o:t these characters it resembl.es in a most striking manner~ h• s.,.ti;o»eata. of central and southern Mexi.co. A large series of specimens froni N$'W' MeXtco and sou.thwastem Texas agree well with those tram Oalitorni.a in the general dorsal.oolo:r pattern (when evident). They ditfer in being sma.ller (max., snout-vent recorded is ,S,'.3.6 mm.; av. 47 .4 mm.)• having a more l"Ounded and longer spade (its length about equal. to its width) and having somewhat larger and more numerous dorsal tubeX"Cles which .are more concentrated dorsou laterally. The tympan'Ulll :Ls small (its diameter ia equal to or less than the distance .from the.nostril to the eye), the internarial distance ;ts slightly- less, vary1%lg irom 2.6 mm. to 3.9 mm. (av. , .. 2 mm.), and the I interorb1tal width is not as great ( it varies .from :3.,2 mm. to 4 .. 7 mm.; av. 4.1 mm.). The parotoid ie faintly evident and the spade reaches the posterior marginot the eye, Indiv.\.duals from. southMatern .Arizona· dif'ter slightly from those •' , -,. . . ' ' ' . ' . . ' . " just disousaed in p~senting a mo)';"e coarsely tubel"Qllla.>:' appearance (1:1e~ pl. IV, tig Tha larger• mo~ numerous tubercles are closely packed .5)•. . . ' . . . . dorsolate?,"a.lly, and the spa~ reaches the anterior Illa.I'gin of the eye.

In all other :respects they are similar. to specimens f'rom Nw Mexlco and Texas •.

Specimens from the northern part of' th~. Mexican :Plateau (b-o.rn .the '· states or ~oahuila, Sonora, and (northel"'ll) Durango) agree well 1tT,lth those studied from the southern.United States. Omt specimen,. however, i'rom

4 mi. W of Saltillo, 5000 :rt.,' . Coahuila. ' (EST-EMS. no!' 2438) 1 ha$ onl:y a faint 1ndioat1on of dorsal tuber~les, giv:lng the appearance of neQJ'ly smooth sldn.,. In all other details it agree~ with th~ others~ Intergradation with mwrooJldl ~JJ.ca;t! seemingly .ooours in southern Dw:-ango and. northern Za~taeas. b"p!&oimens £.rom. th~se areas appear to be intermediate between ht hammon~ and §.~ h~ m.,ultipJ,;teat:a in the degree of dorsal rugosity, size 0£ the parotoid gland, and the l$ngth of the femur and tibia rala.tive to th, length

0£ the body"

It should be -,mpha.sized at this point that orcy average trends are being discussed here and that indiv.1.duals and sometimes entire local Populations may deviate ooneiderably trom .one or more or these trends~ General gimaj,b;gti9n1. . h• oommomi,. ranges from western

Oalii"ornia soutbwestwai-d through Arizona, Ne.1 Mex:too,,. wtllstern Texas into northwestern Oklahoma., n<.>rlht-lard. through wesuarii. Colorado into " . northwestern wYom!ng. Ac,oord:1.ng to Tanner (1941:88) t.bia spadefoot. is also found in northeastern wa.shingto:n and nortlmeste;"'A Montana. Soaroity oi' specilliens .from the la.st thx-ea ~tates,' ano:pauoitj, of literatu:tre records trom this ~ea.··make·.. it ·1mpossib16 to. d? •mo~ .than·· to present a tentative statement ot tjte dir:rt.ribu.tion in the~ st.ates .. ' ' . ' •, .• . - ~. . •, In ~.exico* this specie,s qoours in noJ:"'tb~ Baja Oali£ornia,, ;SOnorat Chihuahua, Coahuila, Nuevo Leon, and. Tamaulipas and rangt;1s southward on the Mextcan Plateau to about La.t:e 2:'.1°N where intergradatio~ trl.th a,. h• ~ip]zi.oati, oeo~s~

Specim!M examined: 3.30; J\!ji.~9.nai Don Lu.is, neal' Bisbee (4 Ktl), .. near Don Luis (23 KU), 1:3.1 mt. E of El.frida1 Coob:i.sa Co. (2 KU); Rad Lake, Navajo Co. (15 FAS); northeast of Tucson, Romero Canon, San~ Catalina Mts .. (1 KU), south ot Tuoson, £lat near Santa Rita Mts •• rPima . .

0011 (1 KU); 1.5 mt.Sot Fort HuaChuoa, Cocbisa co. (1 UI); i mi. ~

Tuoumcari (27 UI) 11 805 Grand Ave., Nogales, Santa Cruz Co .. (12 UI}J

CaJ,fornig1.1 Fa.tr1ngton, Mono take, 6800 ft. (2 MVZ), Benton• Sp00 ft,., (1 MVZ), Conviot Creek, Lone Valley, 7980 .tt. (1 MVZ), S ml.. Wand 4 mi. N 0£ Benton, 6800 :f't,., Mono co. (2 MVZ); Sierra Madre, 1000 .tt. (6 MVZ), 1½ 1111. NW' ot Claremont (3 MV'Z); Century Blvd. and Western AVEt .. ·,, Los Angeles, tos Angeles co. (1 MVZ); QRlors.doa oatet,ray, 4600 tt., Mesa Co. (1 KU) J li,m [email protected] S m:t. N of Glenwood, Ga.t.ron Co. (62 KU)i

Clovis,. Curry Co. (1 KOH, 10 mt. SE ot Silver Oity11 Grant Co. (3 KU); . near Florida, Luna Oo. (9 KU)1 near Insari.ption Rock (37 KU), 1.5 n4• E 71.

ot Grants, Valencia Co., (4 KU); OJ4@b2!¥U .A:tva, Woods Oo. (1 KU)J

Tess: Alpine, Brewster Co. (70 IOJ)J MEXICO: coabui]A.: 7 mi. S and 4 mi. E of Bella Um.on, 7200 ft. (2 KO)J :Parra.$, ,54?0 tt. {'i Iro); near S'inmll1.t. 0£ Mt;. zapalinam6, Sa.J.tUlo, 7800 ft. (12 EHT•HMS); 4 mi ..

W of saltUlo, .5000 rt. (1 EHT•BMS) J ~m,go: 15 mi, Sif ot Cu.enoa.me (1 UI}J Hacienda AtotonilOOt 12 mi. SE Ye~ban:ts, 6680 ft. (6 KU}; /: near Pasaje (1 UIJ 1 EH'l'-HMS)f ®nor,5u 29.5 mi. s ot Nogales (1 UI)J Z,,,aoate9a§.# . 3 w.. S o.t" Avalos (:i mr ...100)1 2 tn:1. N of San Tiburoio,. ciroa 9000 ft. (2 EBT•BMS) •

bimmon©r lIDf!. t;.P+:3,caia (Cope) (pl. r, :r1g. 6j p1. IV, tigs. 1, 2, 3J fig .. 4) ,a(qaphiow) proJ.tipl;tcatU§ Cope, Proo. Aoad. Nat. Soi. Philadelphia,

March, 186.'.3, P• .52. 'fyp$ locality: Valley of Mextco; Brooohi,

Mission Scientifique au Me:d.que et dans l.t Am(riqUE) Centl"3.le;

Etudessur lea Batra.eiens, Liv.i:. 1, 1881, P• 2.5 (listed);

Boulenger, Catalogue of the Batra.obi.a SaJientia in the British Museum, 2nd e., 1882, p. 4,36 (listed); Garman, Bull. Essex Inst.,

vol. 161 1884. p. l46 {listed}; OQpe, Bull. u. $. Na.t. Mus .. , no • .32, 1887, P• 12 (listed)J Gunther, BioJ,.ogta. Oentrali-Ameriaana,

Reptilia and Batracbia11 1901• p. 260 (listed.)J Gad.ow, Amphibia

and ReptU$s, 192;, p .. 164 (general disQUSsion); Nieden, Dae

Tierreioh, Lief. 46J .Anura I, 192:3, P• 48; Taylor, Univ. Kansas

Soi. Bull.,, vol. 28, no. :3, 1942, P• 421 pl. 2, £1.go. 3, .3a, :31,; pl. 3, fig. :3 (tadpole described and £igured)1 Taylor and smith Proo. u. S. Nat. Muse, vol. 9.5, 194.5, P• .551 (State of M6xioo, Mexioo); Smith and Taylor, Bull. u. s. Nat. Mus., no. 194, 1948, P• :;6 ($1ll1'11JlSl7 of distribution in Mexico); smith 8lld Taylor. Univ. Kansas so:l. Bull•• vol. 33, no. 8, 19.50, P• 329 (type locality . ' restl"icted to Ooyoaoan, D. F • • Me.. 'dco); T~1or, Univ• Kansas Sei.

. . .. / . Bull., vol. 3.5t no. 2, 19.53, P• 1.591 (San Luis $tosi.H Orton,

Copeiat no. 21 19.54, P• 98 (dis01,1sses dimorphism in la:rval mouthparts) •

m;u1t4,PJ,icata. Cope, Jour. Aoad. Nat. Soi. Pbila.delpbia, ser. 2. vol .• 6, 1866, PP• 81 ... 821 Cope• Bill. u. s. Nat. Mus., no. it 1875, P• 31 (listed); Yarrmt, Bull. u. s. Nat. Mus., no. ?4• i883, P• 177 (listed)J Cope• tr. a. Nat. Mus., Bull. no. '4, 1889, P• ,308, tig. 79J Taylor, Univ. Kansas sci. Bull., vol. 34, no. 1),

/ . 19.52, P• 798 (six localities in San Lu.1.$ Potosi• Mex:tco)J Davis and Smith, Herpetologita, vol. 8, no •. 4, 195.:3t· P• 14.5. (first

reoord. .from Morelos;.- sexual·. l· diftsrenoe$, . not~d.).'< Soaphio'P'l!fl hoJ.Iy:golq4.. Dugea, La.Natural.eza,vol. 1, 1869 (1869 ...1870),

P• 144 (from "Oua.najuato and oue.dalaja.:ra.0 ). aoapb,iopus s!nm~sU, Broocbi, 13ull. SoQ. Pbilomen., ser. 7, vol. 3, no. 1, 1879 (1878-1879)~ 'P• 23. Type 1ooalit11 "Meld.co"J Broochi, Mission Soienti.f'iqu.e au Mextqua et dans l•.Ame:rique .CentraleJ ' . ,. ' Etudos su:r lea Batra.Qj,,ens, ld.vr. 1, 1881, P• 24, pl~.9, tigs. 4a• 4b (Guanajuato)a Boulenger• Oatalogu.$ .ot •the Batraohia

Salientia 1n the British Mus$um• 2nd ed., 1882, P• 4361 G~.man• Bull. Essex Inst.• vol. 16, 1884, p., 46 (lietedh Duges. La Naturaleza, (2), 2, Cu.ado 10-11, 1896 (1892-1896), P• 482 (known in Mexioo from tlMextoo, Guadalajara, Silao, Guana;iuato"); Mocqua:rd, Bull. Soa. Philoman., ser. 9, vol. 1, p.1t8 (Jalisco)J Lampe, Jahrb. Nassau Ver. Na.turk., vol. SS; 1902, P• ,$1 ("~") l Gad.0"11, Proa•

Z~ol. Soc. London, 190511 i,. 193, 205 (Totol~pan, Oaxaca., Mexico). S2anhiPM hgmmgng:j. l?f9ltipli9atµs, Kellogg, Bull. u. S~ Nat. Mus •• no. 160, ·1 932, pp. 22-24 (distribut1on "in Me:.id.co) (part.)J CChao Liu,

Proo; Boi,ton Soc.· Nat. Rist,., vol,. 41• no. :;1 1935, ·P• ,;O (vocal sao de.aaribed)j Mart:61 del Campo• Anal. Inst. Biol.. ,, vol. 7, nos. 2-3, 1936,, Pi 273 (Hidalgo, M~oo}f ~, vol. 8• nos. 1.2, 1937t p·~ 260 (Vall$ del Mezqtiital, Hidalgo• Mexl.oo)J $mi.th and

La.ute 11 'Trans. Kansas Acad .. Soi~, vol. 48·, no·• .3t 1945. P• :,26 (Mexioo (city), Mexioo}. ~- hammondi mu'.l,tip'.14icata, Firsohein; Herpetologi~a, vol. 6• 19.50• PP• 76-77 (brief discussion of variation). xaxonom;to bke,torj:a .Cope (1863:52) described. Sogphiopus mµ).ti}2liog,- .. tu,R .il;"Om a single specimen aoliected by John Potts {U. s. Nat. Mus. no .. ;3694)iiin the "Valley of MeJd.ao".. For many- yecll9s tl:ie- type was ·the· only

kn~ o:t BJ;>ecimen' the species~ Thia' speoies'was assigned to the Genus . . . by Oo~ (1866.:81)', but fem wo,:,,kars have followed this move. Smith and Taylor 1(19,50(329) ~etrioted the type, looilit.y of aeaphioP,.US mp;l.tiplicatu; 'to (fo;roa~, Distrl.to ,aderal, Mexico. Soappiopus dAAesM, described by Broeobi (1879123) ori the basis of•two young specimens from

"Mmoo" was placed into the eynonyray o.t hammondi. haplmongi l:)y Cope

(1889f30.5)J Kellogg (1932bt22) oorisidered dµgestj. a synonym-of {$cap};4opmt hgopdi pn;lt;t.plioatua.

The status of nmJj;;tpligata has long: been one of dispute. Kellogg 74.

(loe,. cit.) treated it as a subspecies o.:C .:aoa.phio,wf[ hagnthto warty~ separable into at least·two groups•••"• l)iagpord,s: A .~·m..th. closely' spaced la'rge dor$al tubercles,, thase more conoentrated dorsolaterall,y; spad.e rounded, ttSlla.ll3 as wide as 1ong; parotoid gland well developed, often eneroaahing upon tympanum, which is usually indtst:tnct;. spade reaching to between tympanum and anterior margin of' eye.

Va.rj,at1om A large series of spe¢1.meI1EJ from near Victoria de

Durango, sout.i'l$;m Durar.go and £rom localities in northern Zac;,ateQas are generally'more rugose do~sally and laterally than individual.s from looal1ties in northern Mmco, but 119ss rugoae than speeimeM from adjacent Jalisoo and Aguascalientes. Thus it appe!\rS that the aotual area 0£ 1.ntergra.dation between hamm2!¥Y. and llllllt;\,Plicata seems to ba in southern Durango and northern Zacataoaa.

Firaohein (~. oit.) believed the area o:t inwrgradation extended. "probablyi'rom New Me:d..oo to Dur&1go- Mextoo.0 Re adds, "specimens fl"om. this zone a.re dit.fi()Ult to aosign detinitelyto either ~oe but a diatinot north-south gradation may b$ delineated." As noted prev-louszy- (under·~ ho hffi!FU9pg!,) · only average trends are disouesed·hero and oooaaionally entire local populations·may deviate conside;rably ;f."rom·these trends • .A., a.n example ot tb1$ the following notes a.ra presented. A series of 24 speoimena from La 0olo,rada, ·9000 ft., northern Zaoateoas, exhibit a fine i1lU.$tration 0£ the degee,ot variation wbioh mq occur in a loeal populatiollc; ~ha dorsal tubercles vary from tw and small in some indiv.idua.ls to numerous somewhat larger tuberc;las in othe:rs. rn eaob .indS.Vidual th$ do:rf;;ola.te~alwarta a.l"e slightly larger'~ more eonee~rated than those on 'the d.orsum. The small parotoid does not enoroaeh upon· the tympanum.. ot tour speci:alens ·from A~aro, 6400 ft,.• northern Gu.anajuato, tlfO males (EBT.. III-1$ no. 2422.. 2,:n are coar.aezy rugoso like, typi~al .. mu.ltil2la1,oa.ta. and two females (Em' ... m,f.S nos. 242.5~26) have rel.ati vel.y . . . few' and small tubercles. Tbis ·small sample wou,ld suggest sexual. dimorphism, but ths degree of :rugosit;srwash'?t found to be correlated with sex 1n the oouttse ot this study•.

There appears to be, however, a co:ttt-eiation between degree o:t doraal and lateral rugosity and· ei~vatiori. .. Specimens from near Yahtia:U:ca.. 6600 .:rt., Jalisoo (KIJ 110s. 290.$5~.5). Tres CU.-nbrasj 10.500 . ft•• Morelos (TCWM rio. 61~). 19 l!li. w 0£ PElrote, 8000 !'t. (mrr-m,m no. 2.537) and Las Vigaa, 8500 :f't.,. Veracruz (KU no. 26109), and Tenai,go, 9400 f't., state of' Mt::doo (TCWM nos. 10912... 31) (see pl. IV, fig. ·2) • differ generally trom spad.efoots ··from lower elevations in being more rugose, possessing a larger parotoid ( the posterior margin •• of ·the tympanum often being .hidden· ey •it) t Md having shorter lu.nd legs, the spade eoa.rcel;r teaching the.pooterio,.. margin 0£ the eyo. A small series or speo:i.lnenrJ :f'rom .~ ·ws Potos(, QUere'taro* and Veracruz are unusual 1n that they have relativ~:cy smooth skin, a distinct tympa.num1 and the spade is as wide as long. .All are dark and no oolor pattern is evidEmt1 except in one apecimen trom Las Vigas, 8100 ft., Veraoruz (KU no. 26110), which baa two light oval spots on the dorsum just posterior to the eyes. Dr. E. H. Taylor informs me (~sonal communiQation) that he has collected spadefoots at high elevations in VeraCl'UZ whiob wer. very dark in life. Study of additional material from critical looalities within the above mentioned states mq result in a. subspeoific designation tor these dark, nearq emooth-skinned spadetoot toads. Individuals .from the southern limits of the range of ·~ hammonc.li. nmJ,tiplloata, in Oaxaca (UMMZ nos. 11280:3-07 .from near Oaxaca de Juarez) (see pl. IV, fig. 1}, have numerous $om.ewha.t en• larged tubaroles 11 but are not as rugose as speei.msns from Jalisoo, Michoacfui, and Guerrero. In addition the Oaxaca specimens have a shorter bind leg I the spa.de ;reaobing only to the posterior margin ot the eye (spade to between eye and noetril in h• mµltiplieata f'rom northern areas). ThEJ aforementioned epadef'oots £rom Oaxaca. were oolleoted recently and show· taint indications ot oolot". The center of each dorsal. tubercle is light pink. Thia oolor is more distinot on the larger, more numerous do:rsolateral tubercles. The two distinct supraanal spots mu.st. also be pink or red 1n lite.

Laok o:£ sufficient comparative material from numerous areas 1n Mexico is probably tho major reason for the taxonomi¢ instability which has prevailed conet)rning the status ot nrnJ.timJ.cata. In addition to preserved specimens, skeletons·oz )lannnopdi trom

New Mexico and rnµltiplica,:t& .f'rom diti'erent looali.ties in central and 77 ..

southf;trnMexieo have been examined. No Qonaistent osteologioal characters by whioh the two may be se~ted. war& noted. Zweifel (m l4itt.) in:f'ornui; me that he was unable to detect any signifioant ' ' . . : . ' -. . . di:f'ferenoee between Mexican.and Californian specimen$ with respeot . , ,, . ' ' . . . \ ( , to thll skulls. This evidence, Em.pported by the existence. of' a definite . north-south cline leads me to. toga.rd. pn;ltipl;io{i;t8 a subspecies of~ . . . . ; •' ' .. ,, . . ' - ' . hammondi.

in· 0£ However,. . . ' the. . . question: o:f' certain' ' difterenoer:t. . . the. mouthparta. tadpoles of the two spa.defoots presents itself'. Tadpoles multinlicata I • • • ot were first. desorl.bed and figured by Ta:ylor (1942a.:42). He noted that his tadpoles from the state o£ M~co dil"!ere.d from. the tadpoles:\'.-'•.<: . . ..·. ,· ' . . . . . ot pa.mmops!i, described by Smith (19:34:4~9; described. under. thenameo:t·

,soaphioAA bmn,b,.fx:gns_). On this baa,is Taylor :t"Qgarded P!\!1tiplica.ta as being specitica.l.ly diati~ct from hanmi.onsM, •.

The following ·disQUSsion may be a poss~ble explanation for·thEt dii'terences in mouthparts of the two .torms. A e1omparison o:t the figure . ' , .· . ·, ' . . : ·. . ' ' . ·. of mynntondi (Smith, 1934,:pl. 12, :fig. 1) with that of: mul.:Y,pl:tca,t!,

(Tqlo:r. 1942a:p1. :,. £:1.g,. :;) shows that, ·only' slight ditterehces obtain between the two; both possess the diatinOtive median diastema. in the papillae ot the uppe:r lip. It might be mentioned that. this gap 1s not evident in the mout.hparts ot h@.."Wlong! ·:f'igured by. Storer (192.5(39, .tig. EE). In a reoent study ot themouthparts of tadpoles of the pammondi group• Turner (19.52i172) concludes that the dif'!'erenoes in mouthparts are ot no taxonomic value,. . and that tadpoles of h9mM+r

Orton (1954), with her extensive knowledge o:f la.rva1 amphibians points out. howeve:t"• tha.t the probl0n1 of buceal dimorphism in larva&

of western spade.foots 0 ia somewhat more complioated than he (Turner) . ,, . '. has indicated• however. and it warrants mu.oh further etµd.y" .. " She adds that la.na1 di:f'.terenoes are not "constant and wholly ebara.cte~... istic of the respective .forms o:r ,W;aphio;eus. as one would.e:xpeot them· to be." She notes,. "the available $v1.denoe ~geats the possibility . •' that in the hanmtond\ group in generai there is an unstable trend toward overdeve1opment of the mouthpartsasa variant chal"a.cter •• C! .. The exl.stenoe 0£ variant$., (indivJ.duah;, not populatio~s) may thus be the possible answer to the dil'ferenaes which ext.st between ' tadpoles. of M,'lltppndi and ~il2£1qata.

Some JnQntion should b& ma.de of .~rtaiii characters o:t sexual

. . ' . dime>~hism noted in the present study• . 1:he sexes di!"fer. in sizEt 1 the malas average 48.;8 mm. shout-vent length, the females average 46.9 ~.· '. )•, . _.- ' ·- .-. . . .· . in sriout-vent 1ength. .. Thi$ slight size ditf'erenoe in sexes of @?ea h• ntµJ,t,tplica;M ts apparent onl.i in Wge samples. Davis and smith (1945:14$) notedtrutt breeding males possess .;,-, more ext.ensive webbing on the bind toot do tho females. This. . . tb41h;·!:.:-;_ character was tound to be quite consieit&nt in ma~rial. examined by me and the sexes eould be .sepa:rated .ct.u:l.te readily on basis. As mentioned eartiet paper. males of spadefoot toads have in th:i~ all. . ·;· . ' the first two :f'illgers arid often the_third oovei-ed with black nuptial exoreseneea• Otten the irmer margin of the ·lower part ot the :tore~b· is also distinctly oornified~ Qeneml: dislr,tbution: .sm& h• mgl.tip:Y,cata rallges .trom about Lat. 2J0 N in Mexico soutmrard to central Oaxaca. 'fo my knowledge this subspecies has·not bean recorded .from the states of Oolima.

Nayarit; and Tlaxcala in Hex,leo. T'ae southernmost locality from whioh specimens have'I been $xam:i.ned by me is 9.3 ttd.. SE of Oa.."'taca de Juarez, in central 0axac:a., Mexico (UMMZ no. 112807). Gad.ow (·190,51193) records this spadef'oot (under the name soaphiow dyge§.¼) :from Totola'pan, (=Totol{pam), Ou..aoa, the southernmost record station known to me. §pegiplen@ et'9W4ned1 7491 MEXIC01 ,Aguascalienteg1· 3 mi. N of Rinoo'n dEt Romos (1 UI) j 1 mi• S ancl 6 mi. E of Rinoo'n de Romos, 6.5.50 ft. (to KU) g 18 llli. M of .Aguaaoall.entes (1 U!)J 13 mi. N of Aguascalientes (:t tr.I); 6 mi. N of Aguascalientes (1 UI); 10 m.i. E or .Aguascalientes (t.UI)g 4 mi. WSW 0£ .4guaso-alientea (1 KU)J 8 mi• S ot Aguascalientes (1 UI)J 8-10 1111. S ot Aguascalientes (4 EBT-HMSH Dlm1,to federal: Mexico .. (city) (1 UI)i Oiudad Ms"xico, Parque dhapultepe~ (4 UMMZ)J 2.,9 mi. &'1 of Villa Obregon* tt?ed?'egal" {19 UMMZ); 4 mi. s or Villa Obregon* "Pedregai•• (2 UMMZ); Lana, near Tlalpan (1 UI)J Durfmgo,a 9.a Ini. N o:f Viotor1.a de Durango (80 UIH ' . i mi. N 0£ Cborro (3 KUH Guanaju.a.tQ,i 4~ ll'd.. ssw ot San Miguel de Allende• b.r ma.in line of rail.road. (1 ON); Irapuato, 5900 ft,.· (1 UI)f · Aoknbaro• 6400 tt. (5 Em'.,.RMS) I Guen:ertis· hear Mexoala, 2.500 .. 3000 ft. (24 ERT~HMS); Tixt:.la Lake, 9-10 m:L, E o:f' Chipanoingo, 48oo ft. (15 EH'r•BMSh 12 mi. S ot Cbilpanoingo (4 EHT•EMS); naQ- Aoahuizotla

(1 EHT-HMS)I Hidalgo: Actopan (.5 UI); .5 mi. SW of Apam, 7750 ft., (l.KUH Jalisgos near Magdalena, 4,500 ft. (20 mrr-mre); Calle Union1 Guadalajara,

,5100 tt. (7.3 KU); 8 mi. N and 4 mi. E ot Yahual.ioa, 6600 tt.., near tepatitlin de Morelos (9 KU); 1 mi.· N and 7 mi. W of Yahualica, 6600 ft._. near Tepa.titlfui de Morelos (S KU)J 7 mi. S and 1 mio E of Iahua.liea,

6600 ft., near Tepatitlan da Morelos (6 KU); 13 mi. S and 1!> mi.. W of

Guadalajara (2 KtT); Zapotiltio, 5100 i't. (102 mT•HMS)J ?:1629:,90 (State):

2.9 mi. N of 0Chalco (12 UMMZ); San Juan Teot1huao,n (9 UMMZ)I 2:3.1 mi.

NE o:t Ciudad M6xlco, 4 mi, W 0£ Tepexpan (7 UMMZ)J Hacienda sa.n Antonio Ometusco, on Hidalgo state line (3 UMMZ); 40 km. N of Ciudad Me°xl.oo ( 9 UMMZ) J 17 km. ESE of M6x:tc:o ( aity) (3 TOWN); 2 • .5 mi. W ot Toluoa

(7 EHT•HMS) J 3½ mi. S of Tena.ngo, 9400 :tt. (20 TCWM) J Miohoa.oani OojU!l19.tl.fui; near Lago de Ohapala, 5000 tt. ( 4 EHT-.HMS)J 8 mi. N and 4 mi, E of Jiquilpan de Juarez, 5100 ft, (9 KU)J 4 mi .. NE 0£ Tarequato, 6600 ft. (1 KU); Uruapan del Progreso, .5300 :tt. (14 EHT-HMS) J M2rel9~a

4 km. N ot Tres Cum.bras, 10!,00 tt. (1 TCWM) J Progreso, 5000 ft. (2 TCWM) J 6 mio NE of Ouernava.oa (4 EHT-mm); :3 mt.Not Cu.ernavaoa, 5100 ft.

(2 Efrl'•mtsh 2 mi. N ot Cuernavaoa (1 EHT-BMS) J 1.2 mi. N o~ Cuernava.oa

C:3 EHT-HMS) J 1.5 km E o-£ euernavaca (2 EHT-HMS) J 3 km W ot Zaoatepe~, 29.50 f't. (1 TCWM)J 1½ mi. SE 0£ HUitzaJ.a.o (1 TCNlM)i 2 km. s ot Jonaoatepeeo 4,500 ft. (2 T0WM)J Alpuyeoa, 3.500 :rt,., (:3 TClm); 3 mi. S o:t Alpuyeoa• 3500 rt. (1 TOWM); road from Zaoatepeo to Alpuyeoa (2 TOWM); Oaxac§: 7 mi. S of Tama.zulapam (1 UI)I 29 mt. s of' Tamazul(pam (2 UI)# near Oaxaca de Juarez, ,5000 tt. (10 ERT•miS; 5 UI}; 1.7. mi. SE'rof Oiudad Oaxaca, .5100 .ft. (2 UMMZ)J 3.3 mi. SE ot Qiu.dad Oaxaea (1 UMMZH 9.3 mi•

SE ot Ciudad Oaxaca, .5100 :rt. (4 UMMZ); Pu.eblat Oriental (2 EHT•BMS}j

5 mi. E of El Seoo, 8100 :ft. (16 EHT•HMS)J 6 mi. E of' El Seoo (9 UI); 81. near Oumbre (4 EElT-HMSJ 3 UI)J San Martin Texmeluean (1 m:h 3 mi. W and 1 mi. S of tmlfoar d$ Matamoros• 4400 tt. (1 KU); 6.,5 mi. SIT ot Izlfcar de Matamoros (j KU)J 1 mt. N of Tehuitzingo (2 UI)J Qµerjtarqs

Queretaro (oity); 61§0 ft. (12 UMMZ)J 12.mi. NW o:t San Juan del Bio

(2 KU)J .~ ~. f,otoflif Charoas (.5 UMMZ)t 27 mi., NW' •Of Ciud.ad. del

Maiz, Ranoho Tepeya.c, on Mex.le$ll: route 170 (1 CN)l 8 mi. E of Oiudad del Maiz (1 ut); san Diego (9 UMMZ); .1 mi •. mof Sol~ Diez Gut16rrez (10 UI); 20 tni. Nm of San b:iis Potosi (1 UI) J 19 mi. NE of San Luis Pot_osi (13 KU); 9 mi. E ot Ciudad de Valles (1 UI); :rarna.wJ.P,a§,t Miquihua.na, Tampico (1 UMMZ)J Jeru;::u& t El Potrero (2 UMMZ); 2•.'.3 mi. E or Acultzingo (2 UMMZH 1 km. W ot Las 't/igaa, 8,500 ft. (2 KU); Las

Vigas, 8500 ft. (1 KU)l 'l"apeyahual.co, .19 mi. W ot Peroto (1 EHT-HMS; 24 UI); 21 km. N of Perote, 8000 .tt. (1 KU); Perot& (11 ,UMMZ)J zgoat~oa§t 2 mi. S of Majome.• circa 9000 ft. (6 EHT•RMS); La Qolorada, 9000 ft. 24 EBT•llMS); 10 mi. N ot Vtlla de Cos. 6800 tt. (,3 mrr-mlJS); S mi. NW' of Za.ca.tecas, 7600 ft. (1 KU)J 2 mi. ESE of Troncoso, 7000 tt. (2 KUH

8 mi. SE ot zaoate0as 11 7225 ft. (2 KU).

1:ntmtonta.na (Oope) (p1·6 V; f'ig. 1; £1g. ,5)

Soaphiows ogugW,.. Yarrow, aurv, w 100th Merid., 5. 187.5, P• .526 (Pl90vo.

Utah)J Yarrcn-1, Bull;. u., s. Nat. Mus., no. 24, 188311 P• 177 (distr:1.- bution) (~.); Garman, Bull. Essex Inst .. , vol. 16. 1884, p;; 46 scaphiomis :9ou9!}! varj.us, Yarrow, Surv. W l.Ooth M&rid., 511 187,5, P• 526 (Provo, Ut&'l) i Garman, . Bull. Essex Inst •• ·vol. 16, 1884• P• 46 ). 82.

Soaphiopus inter;ontanus Cope, Proo. Aoad• Nat. soi. Phila.delphia, 1883, P• 1,5. Type lc,ca.J.itya Salt Lake City, Utah and Pyramid Lake, Nevada; Ga3:'ll1an;. Bull. Eissex Inst •• -v?l• 16;: 1884,. P• 116

(listed); Tanner, Great Ba.sin,Nat. 9 vol. 1, no., 1, 1939, PP•._ 13-16, pl .. 1, figs. 11.. 12; _pl. J (critical ta:xonomio_ study; tadpole described)J Tanner, Great Basin Nat., vol. 1; nos. :;...4,

1940, p. 139 (Utah); Tanner, Great Basin Nat. 0 vol .. 2, no. 2.

1941 1 P• 88 . ( distribution ill-IdahoH Rodeek, Um:v. Colorado

Mus. Lea.fl.et no. 2, 194)1 p. 4 (Colorado hypothetical list)g Stejneger and Barbour• Cheak: ti.st o.t North Amerioan Amphibians and Reptiles • .5th ed., 194:3,/ P• 39; Ba."tter, Wyoming Wildll!e, vol. 11, no. 8, 1947, P• 3.3 (distribution in Wyoming); Piokwell,

Amphibians and Reptiles o:l the Paoi£ic ·states, 1947, PP• 14, 169, (general discussion}; Orton, Copeia, •no• 2, 19S4. PP• 97-98 ( dimorphism in larval mouthparte disouased.) .. ·

-~ ha-rnmon#, inte;:mon:tamus Cope. Bull. Uo s. Nat. Mus •• no. 34, 1889, P• 304; fig. 76J Oope, Aoad, Nat. Sci .. Philadelphia, vol.,

4.5, 1894 (1893), P• 182 (British Coii.w.bia• Canad.ah intermontan&s Oope, Amer. Nat., volo 30, 1896, P• 101.5 (geographic distribution). aoaphiopus haxn;uS?nc'lj,• Dickerson; The Frog Book, 1906,:;P• .59 {footnote, ~.); Taylor, Univ. California Publae Zool., vol, 7, no. 10, 1912, P• 34.5 (s'l1tm\'IB1'7 or distr.tbuttori)J Ruthven and Gaig~, ocoas. Papers Muse Zool. Univ. Michigan, no. 8, 191.5, P• 1,5 (Nevad.a)J Van Denburgh and Slevin, Proc. Oallf'ornia Aoad. s01., aer. 4, vol. 5, 191.5, p. 102 (Utah); Dioa, Univ. California Fublse Zool.. , · · vol. 16, no. 17, 1916, p. 300 (southeastern Washington)J Stejneger and Barbour, Ohaok List of North American Ampbibiens and Reptiles, 1917, p .. 2!; (part.); Van Denburgh and slevili, Proo. Oalitornia

Acad. Soi., ser, 4, vol. 11• 1921, P• 29 (Utah reoords); Van Denburgh and Slevin• Proo. Calif'ornia Aaad. sa1., -ser. 4• vol. 11, 1921, P• 41 (Idaho records); Nied.en, Das Tferreich• Liet. 46J Anura I, 192.3, P• 47 (~. lJ $tejneger and Barbour, Gheok IA.at of North Junerloari. Amphibians and Raptilss, 2nd ed11, 192.:31 P• 23

19279' P• SS (Utah); Tmmer, Utah Acad. :So1. 1 21st meeting, 1927, PP• 6.7 (U-t.ah localities .llst~)J Slevin, oooas. Ps.pe:ra Cal.1£- ornia Acad. Soi., no. 16, 1928, P• 84, pl. 11, £1g~ 2 (general · discussion) ~.); Tanner, Cope:ta 11 no. 1?1, 1929, .p. ,50 (Utah); Tanner, Copeia; no. 2; 1930, p .. 42 (Utah)J Ta.im.eJ.1', Proc., Utah

Aoad. 'sai.;., vol. 8; 1931• P• 171, pl. i1 (diattibution 1n Uta.hH Logier, Trans. Ca.:nadian Inst. Toronto, vol. 18, 1932* .p. :,19

(distribution in British Oolumbia, Canada); Stejneger and Barbour;· Oheok Li.at of North .Am&rl.ca.n Amphibiatu; and Reptiles, ;rd ed.,

1933; P• 26 Cm!d-)J sv.tlil.a. and SVibl.a~ Copeia., no. J, 1933; · PP• 12.5 ... 126 (Washington) J Wright and Wright, Handbook ot F;rogs · ·· and Toads, 19:33, pp~ 40...41 ~.); Eaton; Btill. Ra.inbow Bridge·

Monument Valley Er.pad., no. 31 1935; P• 4 (Arizona9 Utahh \food.~· Copeia, no. 2, l935, pp. 100 ... 102 {Baja Calif(>rnia, Arlzona.;·Utah) (,pm.)J Cowan, Repcrt Prov. Mus., 1936, P• K:!.9 (distribution in British Columbia, Canada); Cowles and Bogert, IIerpetologioa, vol,. 1, 84.

no. 2, 1936• P• 3.5 (Utah)J Linsd.ale, Amer. Midl. Na-t.; vo. 191 1938, pp. 20-2,3 (breeding habits in Nevada); Gordon• ONgon State Monogr. Zool., no. 1, 19)9. p • .59, fig. 8 (distribution in Oregon)& Graf, Jewett, and Gordon, Copeia, no. 2• 19:39, P• 102 (reool:"ds from Oregon); stejneger and Barbour, Check List ot North American Alllpbibians and. Reptiles, 4th ed.• 1939, P• 29

(part.); Linsdale1 Proo~ Amer. Aoad.. Arts Soi., vol. 73t no. 8, 1940, p. 198 (distribution in Nevada)& Sla.tlirt ocoa.s. Papers Dept. Biol., College or Puget Sound, no. 14, 1941, P• 86 (dist:ri'Qution in Id.aho)J Carl, Copeia, no. 2, 1942, P• l.29 ( (British Columbia, Canada); Oa.rl, Prov. Mus. British Columbia, Handbook no. 2t 2nd ed., 1950, P• 40 (distribution in British Columbia, Canda.); $tebbins, Amphibians of Western North America, 1951, PP• 207-21711 pls. 15, 55 (general disou.ssion) (part.);

Svibla., Copeia, no. :,, 19531 P• 186 (habits in Washington). Sgaphigpµs hapnnondi ha.mm.oust, Englehardt, Copeia, no. 160, 1918, P•

77 (Utah records) i Mills; Herpetologica, vol. 4., 2nd suppl.• 1948, P• S (distribution in British Columbia, Canada).

ScaphioptJ.;h Sn1der, ·Oopeia, no. 86, 19201 PP• 83-84 (Nevada).

3capbi,enµs hapnnongi, wtex,nogtanus • W~ght. and Wright, Handbook o£ Frogs and Toads, 1942, P• 52 (general discussion); EvEmden, Copeia, no. 4, 1946, P• 2.$6 (Idaho); Livezey and Wright, Amer.

Midl. Nat., vol. 'J7, 1947, P• 202 (eggs described); Wright and Wright,· Handbook ot Frogs and Toads, 1949, PP• 120.. 123, p1. 20 (general disoussion)1 Turner, Copeia, no, 3. 19.52, PP• 172-175, .tigs. 1-2 ( discussion ot tadpole mouthparts J mouthparts of no taxonomic value); Woodbury, Herpetologica, vo1. 8• 19.52, P• 44 (habits, lif'e history in Utah); Schmidt, dbeok L1st of North '

American' Amphibians and Reptiles• 6th ed., 19.53, P•: 59; Stebbins,

Amphibians and Reptiles o£'Westem North lunerica, :1.9.54, P• 143,

pls. 11, 13, 17 (general.' account, diistribution)f Logier and Toner.

C:ontrib Roya1 Ontario Mus~ Zool. Paleo.; vo. 41,· 19.55~ PP• 22.23,

(distr.tbu.tion in Ca~); 1 Slater, Oacas~ Papers Dept. Biol., A College o:£ Puget Sound, no. 16; 19.55, p. 137 '(distribution in Waabi:ngton). §eaphO£!,H! bglon5Y.t ;1.ntemonj;gJW!h Chenoweth, Trans. Kansas Aeade Sci., Vt>l. 53, no. 4, 19.50, p.; .5:32 (Utah). I.,.axgngmio M,§torzi This species, described by- Oope (188311.5) under the name of $eap4':qpu.ri 'intermontanus, was later assigned by · him (1889(304) to the Genus~• In the ismne pa.per Co!)9 reduced it·. to wbspeoific rank. Subsequent workers ·have been di:vided in -their treatment oi this spadefoot. Some authors have tollowed Cope in bis subspecific (but not generlo) all.ocation0 while ot~era have regarded. it as a synol:\YDl 0£ ~ndi,e ·Tanner (1939t13), the first to treat the American'spadefoot toa4s ta:xonomicall;r since Cope•s (1889) great worki, oonsidet'ed. intermontana a distinct species~ Stejneger and Barbour placed it into' the synony-,tJy of PAWU12Pr©- in the first £our editions (1917• 192:3, 19:33, 1939) of their oheok list. It was treated as a 1'ull species in their Sth edtion (1943). However, Sobmidt (19.53: 59), in the most· recent oheok list. reduces intemon:fena ·to subspeo:1£ic rank without explanation. He restricts the type locality to· Salt ·'Lal«, City, Utah. Diagno;n,st A~. with slightly EllevaW .tt'Ontoparietal bones; ' interorbital. space tilled with a gland th.at is occasionally lllU.Ch en- J.arged to resemble ,the .boss of bomW;ro1y,n dor$al skin with somewhat , enlarge<'- tubercJ.esJ dorsal color pattern consisting of two dark bars which begin on the upper eyelids, oonverg~ and join just posterior to eyes to torm. a V; .two .broad light:ru.itea be~n on ·postorb:Ltal. reg.ton, gradually e.onverg$ poeterior4r to join on. ode~ (pa:t:-te:rn ·.suggests outline of a harp), two doraola:tex•,a. light lines parallel· the para.verte-

bral linas; a. light line begins .on coccyx, ·courses anterior~ oM third length of bodJr (see pl. v, fig. th spade U$UaJ.l¥ Wider than long; tympanum sznall, distinct; parotoids absent or· :1.ndistinet. )£8.riationc Some indiViduals posseaw a lzy'pertxropbied interorbital

. gland, others meralq have the interorbital. $pace 9n:ty ·sl:1.ght:cy convex.

Fotu:> individuals from Wy'oming• a state :fi'-om which there are· :tew reoo;ids ot ~ntamqntAA?:, have the ntt1oh· enlarged interorbi.tal gland •. Super-. £ioially the resemblance to bomb;tro~ is .most .strlJd.ng. l:n this respeot the miside11tii'ieationa which havl1! :ret.roited in the past between the two spadef'oots are undel'$tandabll!).,. ·Upon raising the interorbital

skin,)hO\-rever, the boss is found to be glandular in ~:p.~on:l;Jma. and and not bo:n;y • The atorement;tonad specimens are darlc and shcm little Ef'lidenoe of a &,r,sa.1 color pa:t,ter.11e · Whet.her this darkened ooncli.tio11, is a nat'llt"al one or due to preservative 1a unknown. Specimens from Idaho and Qregon a.re also ~k. A series of individuals from ll!evada axbibit remarkable 'Ult1.£ortnity in the dietinQtive dorsal: oolor pattemi {see pl. V, fig. 1). When distinct the colo pattern is dia.gnosti~ for the species. anci'tm&£- montana should not be oonf.'w;ad w1 th hanyttondi or 12.omb1froM• a. trequ.@nt ooourrenee in the past. The interorbita1 space is only sligbt:cy:; oonvex in these specimens i"rom· Nevada. Thia condition obta.in$ in ind:1.Viduals exa.'ltl.ned from Idaho and Utah. Too few individuals of ip.,temo~ .trO'lll the llOrthwesterh atates have been available· to me and :c· am· l.Ul9.hle to determine whether or not a correlation between &tribution and· the ~~Pl\T of the inter... orbita1 gland exists. My $mall sample suggests this possibility.

g,~ral &,gtr;tbut1-,cmt' . . . This. speoie$ ranges from sou.th-central. British Oo1umbia in Canada southward th:t'ough e~tern Wasb;tngton and Oregon. and e.."ti;reme eastern. aalif'ornia into Nevada, ,Utah, and extreme northern Arizona.. It also e~s in southern. Idaho. and western Wyoming•

The a.ct'Ual. di.str:1.btlt1.on o:r. :\,p.,te,mon,tana. l1lld th$ preceding speoies• . bgmbifrgns, is still imper£erzym ot ~gi in this paper). Tanner (1941:88)1 howeve~. :tn h13 aumma:r.v- 0£ the hen>eto£auna <>f Idaho p:t"Operly reports tho spadefoot occupying the aout.ber.n part or the State as Soaphiol?J¥L intetmPntanlJ.l.

D,\§oue,§icmt :1.ptemgntana is distinguished £:ram ~. !;tom~ by lacking the boey- :lntero:rbital boss, :reaohing a larger size (max. snout-vent length ,:,ecorded by me is 60,.1 mm. 11 * av. 5:;.1 mm.; in 12.omW;:ona the max. snout-vent. length recorded by me is 57 ,.3 mm.• av• 46.6 mm.). having a sho~r :f'et1'ltlr relative to total length (average o:t

*Tanner (1939:14) ~oport~ a specimen 0£ intermontana wb:loh measures 6Ji.0 mm.; the> average size ot bis sample is .55.2 mm. 114 I .·•,.; I • 49 specimens examined o o~ier specimens / ( I -L ------··-··-··r··49 I I .J-· I ·-·I I -·--·!

-·-· 34

114

Fig. 5 Geographic distribution of Spea intermontana. 88.

ratio TL/F is .. 419J ill J>~. . .• av., rat.io TL/F ts .420}.. having a longer tibia relative to total length (av.. of ratio TL/Tis ~381J in bombi.frons~ a.v. ratio TL/T is .:3.,7), and in having a shorter head length relative to total length (av. ot ratio TL/H is •2.7.5; in bomb.-fro;ns av. o;f ratio TL/H is .285). from ha.mm.ondi JmwmonM,, intermontana di£.fe1"'$ in ha'Ving a. shorter head relativ$ to total. length, a shorter .f'emur relative to tota1 length, a lesser :tnter11arial distance (varies i'rom ,.6 to 4.7 mm.• av n 1s 4.0 mm.; in h• b@mnton~ it varies from 2.6 to 4.2 mm., av.· is 3.2 mm.), a more distinot tymp- am.un, and in having larger. more numerous d.o~sal tubercles. There are also distinct dorsal color pattern dittarenoes between~ a.,pterm,ontana., /J.• h• hanmion£lj,,, and~. bombifrons. z:wei:f'el (1956sJ7) notes that the mating call o:t i1Jm:mo11~ is to~ unlike that of WWJmon.91 and is more reminisqerit of that ot . . pom1?1-trons, The weight of evidence !"a.vo:rs the interpretation 0£ ~ntemonta.na as a Ml. species distinet from h• hamnlondi. Shannon

(195:3:128) suggests that intmon,t§l.1¥l; me;r· be a subspecies ot .~ b,ombi:f.'rPUS• The evidence thus £a.r :Ls inoon61USiVet but Bragg {1945:65) remarks that the tadpole or ]?om2',trow more olosezy- resembles t~t o:t j5.ntermop~ than that o£ MPmJOlJ:

flpeoime§ ~• 7.5; Arl.EiQMt Red l'.,ak<,, Navajo co. (1 F.ASh·

Cal;tform.~1 Deep Springs, Deep Spcl:ngs Valley ,,{4 MVZ). 7 mi •.$ of

Deep Springs p,. o., Deep Springs Valley, Inyo Qo. {.5 MVZ)3 Idaho., Pocatello• Bannoek oo. (1 KU) s Ngyag&: Pyrami.d take, Indian Agency (23 STAN). Th9 Willows; Pyramid Lake• Washoe Co. (31 STAN}; 21 :: , mi. E of Battle 1-it.. 1 Lander Co. (.3 Sl'AN)J 0rngop1 1/8 mi. S of Ma.theur Waterfowl Ref'uge, Harney Co• (2 KlJ)I Utahi 1 mi. N of Cedar City,

.5900 ft., Iron oo. (1 KU)J Wyomi:ng1 14 mi. s and 1 m. W of Kemmerer, 6.550 ft., Lincoln Co. (1 KU)I 10 mi. N o:f Rat Qreek P. o., Niobr~a. co. (1 KU); 30 mi. s of Bitter Creek (1 tu). west side ot Green River,

1 mi. N of Utah border, SWeetwater Co. (1 KO).

PRY.LOOENY The history of·the Pelobatidae dates back.at.least to the Eocene.

Archa@o£8lobates. ~opelobates. and Pglobf¼twps13. are lqlown from the Eocene ot Europe. Noble (1924) desor:1.bsd M,aoropa1oba.tea from the Ollgooene of Mongolia.. Eelobates and a o1osely allied genus,.

Eopaloba!,es.. have been desoribed from tha Lo-.-1er Miocene of Europe.

' . Two genera o:t ~lobat.ids known only as fossils have bean recorded

from North .Axner.toa.: Neosoatjp.omis* -was ciescrtbed by Taylor (1942b:203) from the Upper Pliooene of Kansas and Egn!lobate« was dascrlbed by

Zweifel (19S6i!,) from the Early Oligocene 0£ south Dakota. The oldest known fossil apadei'oot ot North America is Eo®;t.obate; grandis recently described by Zweifel (12£. s.;L!•) .from Early Oligocene beds of South Dakota. He also descri.b9d. (loo. pit.) soaphio12u1 a.lexa.nderj. from the Lower Pliocene of Nevada. T~lor has described

,agapW,opus pliobatraohu§ (19.36), ~• studm, (19.39), and§:• anti93:1us - - . (1941b) £rom Middle Plloaene beds or Kansas. and _a. gtyere and

Neosgaphiopus noblgi (1942b) from the Upper Pliocene of .Kansas. Tihen (19.54) reoords soaphiowg 'gombtt:rons troro. Pleistocene beds in Kansas and Au:f.fenberg (19.56) refers f'ossU material from the Miocene or Florida to Soaphiowg })olbroolg.

*Ritland (19.5.5:44) relegates Neoscapbiopus into the synonymy 0£ ~oaphiopus. 90.

Aooordin.g to. Noble. (ma.. o.·iA.) ttpelobatids a.rose fl"Olll the more primit~ve discoglossid.$ of whichIAot?i~ andAsqaphus are the onl;y surviving representatives." Noble adds• "all evidence points toward a.n origin or pelo'batida in .southern Asia during 'early Tertiary times 11 and to their· ditferentiation, dllring the,Ollgocen$.or Eilar,lier, :.\.nto spadef'oots wbi.Qh have suoc:eeded in spread:ing weH:rtward into Europe and eastward to Mextoo and th&Uxttted state$."

Dunn (192,3s131) states,• 11th& Peloba.tidae are. ma.in4t developed in the l'.ndo-!~ay x-egione bat.. som~. forms. are. found in Europe. and North

Alllerioa,. This is sureli a. cli.spersal b"om a northem oenteit1 the peoull.arity being that· while the Peloba.tirla.e are oonsidt,red a rat.her , primitive f'a.mi:cy, they ba'V'f)·not spread as far.to the south as have the less p~tive bufom.ds a.nci bylida~n

Metcalf {192:31404) attempts to ~a;!J\ diat;-;1.l)ut:ton. oi' saUentia:ns on the basis ot thi!ir opalinid para.sites•· For :Pe1~~t;da.e he suggestEt a.n earliei- or.t.gin, stating the pelobatids ori~ted "in sou1;.heastern ' -,,._., Asia soon a.i.""ter the separa~on ot Australia from A.Sia at the beginning of the Cretaeeous period.,n

The p;oope:t" recognition of." primitive obaraoterais. the only support. tor a valid and useful b;ypothetioal pattern of." pbyiogeq. Noble (mi.. ~-) disousses the origin and evolution of the Pelobatidae and states that "the more generalized pelobatids as x-epresented by H~=uzo:phr,l§

( considered by him the most primitive genus· 1n the, subf'aniil¥ Mego- phryinae) have advanced beyond I,iopelmidae (~bioh a.re. ancestral to Pelobatidae)i.in: (1) the ossification of the illtervertebral cartilages (cantra) which usually adhere to the more anterior vertebrae forming a 91.

procoelous vertebral col'1!ll?lJ· (2).reduetion o:t.the numb(ar of presacral

vertebrae ·to eight; (3) loss of.,;.\cl.W apparentl;r by fusion nth the vertebraea (4) restriction o:t' the ossifica.t1011 ot th& .ventral pa.rt of

soapuloco~coj.d aud the development ot a,-1 acrood.on;·· {.5) redtiotion of

pubis and expansion of sa

ly ear:cy, developed excessive bony or calcareous deposii;s in their dem for we find suoh encrustations not· only in the integument· of the head •., but also along.the baok of aevera.t spaciE.'S of M2goplµey;s." The eeoond stage iu the progressive modi.£1Qa.tion 0£ the pelobatids aocordir,g to Noble (.sm.- ~.) 1a repx'esent~ by thitt ;tossil .lisorowlobates . . ,· - ' ' from. the Oligocene ot Mongoi1.a.. 'i'he important f~a:tU1"8$ gre.·· the .enlarge- ment.· o:C ·the preha.J.l.ux and anky'losif;l ot the bocy ·encrt.1stationa to the akulla due to tossoria.1 llfef .an¢ a gr:,eate:.r e:x:pansion or the aaora,.l .· diapopbyses. Now the Ne-ri' World '.S:AAP~ :and the .Old World £,elobate~ . : , ' .. , ... ·,' . ' ... $hare these :reat~e1:1 with l;laorgpe1o:tm:tes ...· .The t"el.atA.onships 0£ .thes

of pelobatids. The Genus was !ormerly known only from~ speoiss

from late Oligocene and early Miocene beds in Europe. The significant

new fossil pelo'batid, described by Zweifel (1956::3) as E2J2a1obates

grfflldis.. is si m1 l ar in a numbe:r o:t characters to the recent Asian Genw, Meggphtt;§. judging from ·ttu, descll'3.ption. Zweifel • .9i'!i•) points out the OIU.3'" 1t1ajor dif.teren()o that oan be dtscasrned between

Megophrx@ and Eopelo}ates is in the·· greater skull-roofing of the latter. EsrnelQbatea ptobably lies 1n the megophrine line of descent. Zweifel <.9,n. 9!1.) notes th.at tha prj.nlitive anc~stor probably had considerable dermal bone on the skull. Acoo:ding to Zweifel (1956115) the Genus _Eo'P§loba.tes -probabl,y had a holarctic distribution in earl.$ and middle Tel'tiary time. He adds, "Clima.tic Qhange through the Tertiary eliminated F;gpelobate;! from the .American and European faunas and restricted its descend.anti:.

Megoply;':ya to the Asian region where, with the great:cy- increased

mountain-building activity of late, Tart.:J.ary and· Quaternary- t.inle,. the montane genera Squ.tigo;r .. Aelurophr;yne, V'ibrl.ssonhora, and Qpbr;y;ophr;yng evolved." The only evidence fortha assumption that the Pelobatidae arose in southeast Asia (Noble• 1924) is the presen:t ~striotion 0£ primitive palobatids to that :region. ZWeif'el (~. Ja!•) states, "The demonstra• tion of a holarctJ.o dist:t'ibution ot prlmi.tive·pelobatids doea.llOt naoessari:cy- invalida:te the assumption ot Asian origin• but serves to emphasize th& caution.neoessary.when undertald.ng to-•in:t"er·pa~t distribution and dispersal. J'IOUtes i"rom present diatributiona.11 Although EopeJ,obates :posaib4' stands ~ose to .the 'base o:r .the pelobatil19 (spadetoot) line of descent• it 1$ probab1a that the li.ne bad alrea~ been. establishe

Asian Macrg:gelopatss. separation of th& two types ma,y have .occurred· somewhat earlier. ZW'eifel (sm. .. ), as noted above, suggest.$ that. i£. a spade.foot with a well-developed skull· root and·.~. stl"Qng palatine bone can be _ considered prim:L tive fl then Pelobate a a:nd liaorowobates .rep:r:-esent ' the more primitive type and §oa.phj.opuf.!, tbco:ne slightly morer:redueed than

§capW.opus• 8?ld also has a frontoparietal fontanelle, oan on this basis• be considered· more specialized•.. I h.e1-e presont additiona1 evidence to support; the idea ·that. f!eJJ!ba~ and .§.aapb;ppµs, cilfei. ·. tbs more primitive, and~ the more specialized, npadef'oot toads.. In Pelobates and in ~aphl.optlJ! the pl:-ehallux is oomposed 0£ two or more bones and thus is regarded as segmented(~c

ing to Noble the prehallux. ot Maoropeloba.t2s. seama 1;.o be unsegmented. The evidence .:trom paleontology and morphology tavors the inter-

pretation of feJ.oJ:mteta and ~oaphi.opu§ evolving from the same .fossorial stock.

Sc§plJ1oltY§ S2l¼Cbi, · in lacking the segmented. prehallux and having .. . much less skull rooting than ,§.. holgrpol,g and ,a. ·.~is seemingly- the most specialized member <>£ the Genus sca;pM:opu1. Thul _trend of oouoW, toward.~ is also ma.nii"ested in its distribution and habits. j3gaphio1m2 Qouchi is found in xeric habitats,. like all members of~