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Intrageneric Relationships Among Colubrid Snakes of the Genus Geophis Wagler

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Intrageneric Relationships Among Colubrid Snakes of the Genus Geophis Wagler MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 131 Intrageneric Relationships Among Colubrid Snakes of the Genus Geophis Wagler BY FLOYD LESLIE DOWNS College of Wooster, Wooster, Ohio ANN ARBOR MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN JULY 26, 1967 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies bascd principally upon the collections in the Museum. Thcy are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and indi- viduals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, are published separtely. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan. LISTOF MISCELLANEOUSPUBLICATIONS ON REPTILES AND AMPHIBIANS No. The amphibians and reptiles of the Sierra Nevada de Santa Marta, Colom- bia. By ALEXANDERG. RUTHVEN.(1922) 69 pp., 12 pls., 2 figs., 1 map ... $1.00 No. A contribution to a knowledge of the herpetology of a portion of the Savanna Region of Central Peten, Guatemala. By L. C. STUART.(1935) 56 pp., 4 pls., 1 fig., 1 map .......................................... $0.50 No. A contribution to the herpetology of the Isthmus of Tehuantepec. IV. An annotated list of the amphibians and reptiles collected on the Pacific slope during the summer of 1936. By NORMANHARTWEG AND JAMESA. OLIVER.(1940) 31 pp. ..................................... $0.35 No. Studies of Neotropical Colubrinae. VIII. A revision of the genus Dryado- phis Stuart, 1939. By L. C. STUART.(1941) 106 pp., 4 pls., 13 figs., 4maps ............................................................ $1.15 No. A contribution to the knowledge of variation in Opheodrys vernalis (Harlan), with the description of a new subspecies. By ARNOLDB. GROB- MAN. (1941) 38 pp., 2 figs., 1 map .................................... $0.35 No. Taxonomic and geographic comments on Guatemalan salamanders of the genus Oedipus. By L. C. STUART.(1943) 33 pp., 2 pls., 1 map ..... $0.35 No. Home range, homing behavior, and migration in turtles. By FREDR. CAGLE.(1944) 34 pp., 2 pls., 4 figs., 1 map ............................ $0.35 No. The amphibians and reptiles of Alta Verapaz, Guatemala. By L. C. STUART.(1948) 109 pp., 10 figs., 1 map ................................ $1.50 No. Studies of the black swamp snake, Seminatrix pygaea (Cope), with des- criptions of two new subspecies. By HERNDONG. DOWLING.(1950) 38 pp.6 figs., 1 map ............................................... $1.25 No. A brief review of the Guatemalan lizards of the genus Anolis. By L. C. STUART.(1955) 31 pp. .............................................. $0.50 No. The anatomy of the head of Ctenosaura Pectinata (Iguanidae). By THOMASM. OELRICH.(1956) 122 pp., 59 figs. ........................ $1.85 No. The frogs of the hylid genus Phrynohyas Fitzinger, 1843. By WILLIAME. DUELLMAN.(1956) 47 pp., 6 PIS., 10 figs., 4 maps ...................... $0.70 No. Variation and relative growth in the plastral scutes of the turtle Kinoster- non integrum Leconte. By JAMESE. MOSIMANN.(1956) 43 pp., 1 pl., 24 figs. ............................................................ $0.75 (continued on inside back cower) MI IS(:El,I,ANEOUS I'UBI,ICA'I*IONS RiIIJSEUM 01; ZOOI,OC;\', IJNIVERSITY 01; MII(:MlCAN, NO. 131 Intrageneric Relationships Among Colubrid Snakes of the Genus Geophis Wagler ANN ARBOR kIUSE1JbI OF ZOOLOGY, UNIVERSITY OF MICHIGAN JULY 26, 1967 Skull ;~r~tll)cr~titiorr ....................................................... 2.4 'I'llc ltl(.~~tityof the 'l'y1)c Sl)ccic.s. (;c.o/)lli.\ c~lrnlybc.r~t(LV.lglcr) ................... 3fi I)clinitio~~;rlltl C:o~~rl)ositio~rof rlrc (;c.nlrs .................................... 40 IS. \';~ri;~liot~ii~ SI.RIIN.III;II ~OIIII~S ill (;. \~~~ri~loli(ilti\110111 \c\(,I;I~ loc;lliti(.~i ill \'I,I;ICI.II/. hli.\i(o ........................................... I l(i SMAI.I.,SECRK~IVI colt~l~ritl s11;lkes pose special problclns to systematists. Ow- ing to their small siLc ant1 sect.etivc habits, they ;we generally collected only slmri~clic.ally;inti it1 snlall numl~et-s,ant1 cotlseq~~ently;n-c poorly rel~rescntetl in mtlscum collcc.tiolls. Solnc of these snakes possess cllai-acteristics whicll 17-i;lke them, at least st111et-fici;~lly,urliclue; thesc arc often treated -'1s : 1110110- tyj'ic gene]-;{.At (lie ol)l)osite extreme are the large, 1)olytypic genera such as tllc o~.ictit:tl (:trl~r~t~oritr,the Sor~tll/\mcric;tri ,4trtrrt1rs, ant1 the (:entl-:tl ,4111ct-i(.;1tlGcophi.~. \i\fithil~ tl~csc groups, gene flow between populations is ;~l)l);~t'en~lyr;tthcl. rest~.ic.tetl.-1'lle resulting l,ol)~tlationaltliKerences 11;tve let1 to tile clcsc.l.il)tio~lof numcrous no~niti;~lUorms, including some 80 in Cnler- tttnrici ;~ntl,4trcrc.t~r.s ;cntl :tl)out hall' tli;~t11111nbct- in (;c:o/)ltis. linJ'o~-tt~tt;~~cly, thc r;ttlier cotlstal~tinct.ease ill the nrlmber of descrihetl lor~nswas not iicc.onl- ]);~tiietlI)y syt~thcses;~t the generic level. 01ily 1.ecently 11:tvc tllcse syiitlleses I~cc~l;It relnl~tetl. M;irx: xntl Ingcr (1!)55) cliscussetl tlle range of variation ant1 he evolutionary tl.c~~tlsin Crtlcr~trtr~~ic~,:uncl S;tvage (l!)(iO), ;IS well as tlcfining sl~cciesg~.oul)s, (tit1 the same l'or E(.~t;itlot-i;~~lAlr(rc11r.s. :I'lie genus (;ooplris 11;rs beeti in j,at-lic.rllal. nectl of 21 str~tlyol a revisionary tiaturc. Unlike Ccrlnrncrritr arltl Alrcrc./lr.r, which tlespite the 1;u-ge ilumbcr oU ineltitlet1 Corms ;Ire relatively I~olnogcneot~sgl-ottl)~, CGcophis inclutles a highly tlivc~.sc:II.I.:I~ oC snakcs. 7'11is tliversity has leUt Geophis systcrnatics in a cha- otic state; this stutly was ~)rollll~tetlby tl1;tt ch;~os,a11t1 tletlicatetl to t.cclucing it. Rily ~)~-inc.il)lc;tinls 1i;lve been three: (1) :ul ;~nalysisof intraspecific. antl itltt.agctlcric val-i;~tiot~;(2) the establishment ol sl)ecies groups within C~eofiliis wllic.11 (.;In I)c rnea~ii~igfullyt.oml)aretl wit11 one :tnotllcr antl with other gcllet.;i; ant1 (3) tllc synthesis ol cttrl-ent kilowletlgc ol tllc genus illto ;I single volr~llic. M~liatevcl-value this stntly may h;rvc is in large ~neasui-e21 I-eflection of the generous c.oopei-ation of others. For making spec imens in their care ;~r~ailiible to me, ant1 in many ill~ti~llce~foi- examining material at my request, I am gl-;~tefulco SI-. Migllel A1v;rl-ez tlcl 7'oro, J. (:. R;~ttersby,(:lia~-les h/I. Rogcrt, J;llnes E. Uiihlke, Doris R'I. Coclil-;in, \4'illi;1ln 1%. Davis, ,lames R. Ilixon, \ATilli;im F:. I)ucll~n;~n,Josef Eisclt, Alice <;. C:. C;l-;~~itlison,Jean (iuibi., Kobert 1;. Ingcr, :\I;III E. ~,e\lrton,George N.L,owcry, ~retlcl-ick'bletlem, George S. Rtlyr~.s, I<cnlicth S. Nol-ris. <;iinthcr Petel-s, I)o~iglas(;. Robinson, Jay M. S;iv;ige, Hob;irt M. Snlitli, Robert (;. Stel~l~ins,Ernest E. Willi;ims, ;lntl Kichartl G. Zweifcl. In atltlition to tlie ;ibovc, m;rlly pcol~le11;1vc 111-ovit1etli~ilormation and ;~tlvic.ewlric-h have atltlctl iirun~c;rsur:rbly to the toinplctcness ol tllc study Among rllcse wcrc K1.;1ig K. i\tllel., ,\. (:ul-tis ,-\llen, l<i(.hi~rtlE. Ethel-itlgc, Fretlerick K. (+ehlbilc.l~,R/I;lx K. I-Icc-llt, 147;ilter Hclllnic.11, 1)ale L. Hoyt, i\r~ioltl C;. Klugc, Etll~ioi~clV. M;iln;~tc,Gl~iscpl~e S(.ol.tccci, I.;i~ircnc-cC;. Stunl-t, b1enic.o 7'01-thio, ;lilt1 -1'llom;rs R'I. U~~ell.-1'0 ;ill of these 1 give lriy sincc~.et11;111ks. bly wil'c ,jo;tnne tlesel-ves sl)et.ial tlianks. Hcl. many 11o111.s01' typing, conn- Ijinctl wit11 her intercst ;~ritlc~icoril-agelilelit, aitlctl mnteri;rlly i11 the coin. 1)letion of the stirtly. My coilunitccc 1i;rs ;illoxvctl III~to tlevclol, tliis stt~tlyill 111)' oxv11 way, xvI1ilc at (Ire siilrlc time ~,enl;~ir~ilig;~~~;iilnI~le fol. atlvicc ;r~ltlc-ons~iltation. For this 1 ;II~tlcel~ly itl~prcciativc. I am in s1)cci:rl tlebt to Ch;il.les I;. \4';1lker, who at lny rccl~icst ex;inlinetl ty~)(!nl;i(wi;tl in sc\~e1.;11E~iro1)e;in collectiolls, wl~o cllrorlghollt tlie stlltly h;is heen ;I const;ult source 01 ;rtlvic.e, ;~ntlwho, ;it ;I ti~neof rn;~xil~\nltlcm;intl on Iris ellcrgies, gcncrously ;igrced to iiss~uillcthe c-l~ai~.~n;i~~sllil)of niy colnlliittcc. This investigation was l)cg1111,;rntl ill 1;rrgc 1):u.t colnl~lete(l,rintler the tlii-ection or thc late Nol.ln;tn Hal-tweg. His ;rtl\.ic.e ant1 l)l-obing cluestions were inv;rluable assets lo the stritly; to liilr~I am tleeply gl.iilcf111. SNAKES OF GENUS GEOPIIIS 3 MATERIALS AND METHODS This study is basetl on the examin;rtion of some 1610 spetitnens of Geoplzis. Thirty-one of these have been examinetl by others at my request; the remainder I h;rve examined personally.
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