yy, a es ales \ Pa ORES N SION | Sacra _ wre Vk \ Cl Y oy SMITHSONIAN INSTITY UNITED STATES NATIONAL A BuLLeETIN 154 a. ot ae hr oe + ¥, A STUDY OF THE TEIID LIZARDS... Z “To. OF THE GENUS CNEMIDOPHORUS WITH SPECIAL REFERENCE TO THEIR BoYCOGENE TIC RELATIONSHIPS BY CHARLES EE BWR Of Trinity University Waxahachie, Texas UNITED STATES GOVERNMENT PRINTING OF FICE WASHINGTON : 1931 For sale by the Superintendent of Documents ,Washington, D.C. Price 80 cents (paper cover) %) " ae Bese est re an o oe vee ADVERTISEMENT The scientific publications of the National Museum include two series, known, respectively, as Proceedings and Bulletin. The Proceedings, begun in 1878, is intended primarily as a me- dium for the publication of original papers, based on the collections of the National Museum, that set forth newly acquired facts in biology, anthropology, and geology, with descriptions of new forms and revisions of limited groups. Copies of each paper, in pamphlet form, are distributed as published to libraries and scientific organ- izations and to specialists and others interested in the different subjects. The dates at which these separate papers are published are recorded in the table of contents of each of the volumes. The Bulletins, the first of which was issued in 1875, consist of a series of separate publications comprising monographs of large zoo- logical groups and other general systematic treatises (occasionally in several volumes), faunal works, reports of expeditions, catalogues of type-specimens, special collections, and other material of similar nature. The majority of the volumes are octavo in size, but a quarto size has been adopted in a few instances in which large plates were regarded as indispensable. In the Bulletin series appear vol- umes under the heading Contributions from the United States National Hebarium, in octavo form, published by the National Museum since 1902, which contain papers relating to the botanical collections of the Museum. The present work forms No. 154, of the Bulletin series. ALEXANDER WETMORE, Assistant Secretary, Smithsonian Institution. Wasuineron, D. C., Marcy 17, 1931. Jil TABLE OF CONTENTS Page. NG MOCIUICULOT Reina cry es ee ee Ole ee NE ae ere oe ae eS 1 SCO ERC A MECC Vie eee as ee eet meee ee Bene ee ee ee a ee 8 PeEMIEMOTC UNO pROPUG! = tae fa Bes ee Ao eect SSeS 10 (Gommmromemaim essere ee es ee ee ee See ee ee 10 SREBRTLTY IT renee oes ee i eS eee eS eer A ie SE 10 SPECT MGIC RO lest ee els ee ee Se eee ke Ps Ek eS Se oo 11 MORSE ONES Ets Nesey ee Ne eet Oe SAE RL Oe OS en a ee 14 ECE rIptrmi en sce ers Boats Bee ar pee Sy Be ye 15 Rsk TEI Sh Ei Te ne os ee es re ee Se et ee BS ae ee 18 Pp aE me a a ee a ey pe ee ne UE te te ee a St 18 Elst tie Ue ee eee Ree Ne ee ey ee ee ee a eee 19 ATED by 1 i eee eee ew nN Ache 4 Ren OS EAE BE eo ee 19 SLOGiImisionsOnuhe cenus Cnemidophlonusn= === == a= == ae 20 Horms wrongly assisned to Cnemidophorus..__..-.-=--.----------===- 20 Meavahonune forms OlCrvemidophorus..- 2-262. 220. = Sea ee 21 WBMES ANS TOUD Sons Lee eee Se eo oe ee eel as 30 GHILEMMNISCOLUSMIEMINISCOLUS Be ss ae ne a ee ee Se 30 DCT RISCIEUS NUGTUCOLON see teen, eee oS Se 2 hat ae ee ee a 40 GELLING MRA e eee ae ee A eet a ia ae eee EE SR 43 CRAG ETUUS EIN UNLV Sea ee ae Eee eS Se es i eee 46 ORT TILUSRCLTALD C1US US et ee Ee SS SS ee 51 SOUT ENTE Gok a i dh ie ee ee eng eee ee 53 SPUR TA ETO Ne Sete Rye foe es rs Ces Ay oe a 56 LED DIAC D DIU acts Ae een eee ees ae Pe a Ne ee See 56 MELE N DUTUCOUMECLIS: Sie Aa ye ne pene oe eee ees Et 63 Cs STROSS Oe Me SI else sis Raven yt OB nga ay om Rees yok ere oye ee 66 S (RICCTNG ON eS eee ag Meech = lee ts 2M Ra gn Oe eS oa ae a See 74 SR RCreEMMCALURTE TOU eee tO eee ek ee ee 76 (CRESCRIUTICALUSESCLUUTICALUS men ee ee oe ee ee eee 76 RESET TEAL UGNTULOTTS es = hie See ee Sr el Wee SS aa 97 PRS EEECTILUS HON LOL Rope pie ofan hal Bee eee a tee Sey dE 122 (CRE OU See eee ee et ieee oa Uy) | eee 5a BS eee ee ee 141 oer Va salah eee ee See es ee eet Beene to ere Uo Fe hae 144 Meer escell viusserGy oe 26 02 oe oe ee ee Oh eet ee bee 146 Giiiesscllavustiessellatus== = 2 ae = eee ee See Ee eee 146 (Gmtessellatustiuvdusee= ook oa ee ee ee ee eee 199 MAME PESCHLATISRCCHONIEDOR 8 <== 2 = Rae Ee ih ew ome Bee LR ee a2 202 OIESSECLLGLES NONLNIS 22 = Soe Ske en eee ee ee a 205 GMmiCSSCllatuSiCONUSS 2.082 Sat eee ee eee ee ee ee ae 208 (GERUCCAL Saree ne BN Se ee ee ee ee ns eae Ad es ZA @=catalinensis= = =—— = I a a ee a er ne a ae 213 OMCCEOLOCTISTS SEs = te ee Se a ee ee cates 216 GEETLOE TT Se ee eS Soe R22 a Pee Se eS 218 SER ee eeer ene: SF RR Ey cE eh sb ee Lee oe 221 VI TABLE OF CONTENTS che hyperythriusictoupis=.2 fa 25 2k Se. ee ee ee ee ee CRG DEryERM USP ILYy DENY CLETUS rece ee ee a ee ee Cphaperyennus, CACTULCIS === a ee ee se ee ee CSG DEN ERT US DUCLUS ee ee ee Cchyperginnus danherniag 5 ls se eee ae ee ee eee ee er SUMUIMaT yea 22 2 se Pas Oe ee ee ee eee oe eee eee SME CleslOl UMCELbAl ta xXOM OMT CH OSUl TOM ee eee ame ae a Cisachktt 2. 2 2 ot oa ee ee eo ee ee Generale SCusstorano hae) ey tO 10S 11 See ee SUOMI AT ees er ee ee ee Syn lr ae ieey ee Biblioonaphiy's= §- "2.22225 5522 ee SE ee eee ee linden eves = Skee ee eee eee Page. 226 226 240 242 244 246 249 249 251 260 260 283 FIGurRE 1. COST Oo Ore GO DO © 10. Wut 12. 13. 14. 15. 16. ie LIST OF ILLUSTRATIONS Diagram showing tongue characters of typical Cnemidophorus INCU DIC Al EVA ILCTD UO Seeley ee ee PE ONeMIdONLOTUSMLLGLvuS-a) LOD On Mead ==) si. == ee eee . Cnemidophorus tessellatus rubidus. Side of head_____________ . Cnemidophorus tessellatus rubidus. Gular region____________ . Cnemidophorus sexlineatus gularis. Mesoptychial region___— __ . Cnemidophorus lemniscatus lemniscatus. Anal region________~ . Cnemidophorus lemniscatus lemniscatus. Color pattern ______ . Map showing locality records for Cnemidophorus lemniscatus NOUIVESC CGS oe ee ORD oe Oe NG EM Te ee Oe RE cs ee . Map showing locality records for Cnemidophorus lemniscatus TEL OEUCO LOT ee el ee re ee i Map showing locality records for Cnemidophorus ocellifer_____ Map showing distribution of forms of lemniscatus group _____ Diagram of the supposed relationships within the lemniscatus LO UL) ee a oat ae a RINE CI 52 OPS NE we et ES. 5 ea SE Map showing locality records for Cnemidophorus deppii deppii- Comparison of typical color patterns of Cnemidophorus deppii cozumelus and Cnemidophorus deppii deppii_______-______- Cnemidophorus guttatus. Variation in the dorsal striping____ Map showing locality records for Cnemidophorus guttatus_____ A theoretical diagram of the manner in which two closely re- lated species, at present in co-extensive distribution, may have evolved from a common stock through isolation____ ~~ . Map showing distribution of forms of deppii group___________ . Diagram of supposed relationships within the deppii group __- . Map showing locality records for Cnemidophorus sexlineatus CY Ein POLL A OAT a ie es aR Sg eS ae Ls SS Set eee =f iy . Map showing locality records for Cnemidophorus sexlineatus CORY RAG POSS cS a See Nay rae ras Po ee ee fe gL . Cnemidophorus sexlineatus perplecus. Variation in the color PD Skt GET Niro eee: Reena, Srey eee ee Ee . Map showing locality records for Cnemidophorus sexlineatus DICT Ue I ws A 159 es ES NE SD! ae ee NES a a we 2 oe . Diagram of the relationships of Cnemidophorus sexlineatus DOR UCTRLS 5 ere = ess a5 aE EN See eee Soe Le ee ee 25. Map showing distribution of the forms of the sexlineatus group_ 26. A diagram of the supposed relationships within the sexlineatus PTO (Nm es te Oca ees dee . Map showing locality records for Cnemidophorus tessellatus LOSSCLICUILS ee ge Nee ae ee Be ees aeons Se ee Be . Diagram of the supposed relationships of Cnemidophorus tes- REL DIAS COS SCRUBS Es ets See A ee ee eee ee ee Cnemidophorus catalinensis. Color pattern________________- . Map showing the distribution of the forms of the tessellatus OT OU Tae a a ae SS SSeS SS Se bo bk wNonwnbd He Ww ‘S 133 140 145 146 Noe cn VIII FIGURE 35. 36. 37. 38. . Color pattern variation in the hyperythrus group . Map showing distribution of the forms of the hyperythrus LIST OF ILLUSTRATIONS . Diagram of the supposed relationships within the tessellatus group . Map showing locality records for Cnemidophorus hyperythrus hyperythrus STOP: at ao. ae ae Le eee oe a a ee Diagram of the supposed relationships within the hyperythrus OTOP Ae Soe Se = Se aa ee ee pe Diagram of the supposed relationships of the primitive Teiidae_ Diagram of the supposed relationships within the genus Cnémidophonus* 225226 2 ee Map showing distribution of the five groups of the genus Cnemidophorus Page. 225 237 247 248 249 253 A STUDY OF THE TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS, WITH SPECIAL REFERENCE TO THEIR PHYLOGENETIC RELATIONSHIPS By Cuaries E. Burr Of Trinity University, Waxahachie, Texas INTRODUCTION The genus Cnemidophorus, one of the largest and most important units within the family Teiidae, is a complex assemblage of species, subspecies and phases, which reaches its maximum diversity in North America. The forms here assigned to this genus occur from southern Brazil to the northern United States, both on the main- land and on the closer neighboring islands, thus comprising one of the most widespread and characteristic groups of lizards of the New World. Within the remarkably narrow generic limits of Cnemidophorus, an amazing amount of variation is seen. Its unusually extensive distribution brings it into contact with a great variety of habitat conditions to which its forms seem to be constantly adapting them- selves through their apparently natural tendency to vary. More- over, in many of the species each individual shows an intricate series of pattern stages during development, and abnormalities are frequently found in the various populations. Many of the sup- posed species have been described from only a few specimens, often wholly without regard to the geographical probabilities, and fre- quently without reasonable allowances for the normal develop- mental and environmental pattern variation. Furthermore, the genus itself has never been clearly defined and it is found that some of its oldest species should have been referred to the very closely related genus Ameiva. In the absence of a comprehensive review of the genus Cnemidophorus it is not surprising that there has always been much uncertainty as to the number of forms, their relationships, distribution, and differentiation, one from another. The present study has been undertaken to bring order into this perplexing group of lizards, and to discover the origin and relation- 1 2 BULLETIN 154, UNITED STATES NATIONAL MUSEUM ships of the component forms. It is hoped that this revision will stimulate interest in the various lizards of the genus Cnemidophorus, that it will show from what localities specimens are needed, and that above all it will emphasize the lack of knowledge of habitat and habits. The need for this work has been evident for a considerable length of time. Cope (1900), after a lifetime of study, wrote that “the discrimination of the North American species of this genus is the most difficult problem in our herpetology.” Prior to this a foreign viewpoint was expressed by Giinther (1885), who elaborated his dis- cussion of Cnemidophorus sealincatus by the following comment: “ Bocourt included in the synonymy of this species a number of other names which have been created by American authors on very shght grounds. I fully agree with him as to their value, but it is difficult to determine to which species they should be referred. From the character of the descriptions and figures I regard this task as impossible.” The opinion of Gadow (1906) is well expressed by the following extracts from his work on the Mexican species of Cnemidophorus: “This Teiid genus is invaluable for the study of variation. It is so plastic * * * that it is represented by some form in almost every kind of-terraim.” * “ * "Most. of the: “species =are™ce variable that they may well drive the systematist to despair. No two authorities will, nor can, possibly agree en the number of ad- missible species’) * -* * (for) * * *\-almostvevery one/ot the taxonomic characters investigated in this paper has an amplitude of variation within some species which equals that of the whole genus.” Much the same idea was expressed by Ditmars (1907) in prefac- ing his treatment of the genus Cnemidophorus, thus—* In the prep- aration of a resumé of the species of Cnemidophorous, the writer finds himself confronted by the most difficult proposition of any vet encountered in this work. Provided with a fine set of specimens, he has attacked this obstacle from every side—and with very little success * * * Nothing can straighten out this problem but the work of some one who makes a specialty of the genus, gathers about him elaborate series of all of the species and notes his observations impartially.” It is noteworthy that Camp (19160), in reporting on a collection from California, cited Cnemidophorus as a “ remarkably unstable genus,” and that Schmidt (1922), in his consideration of the herpe- iological fauna of Lower California, wrote that “ An adequate under- standing of the relations of the 15 species of Cnemidophorus in the present list can only be reached by the comprehensive study of a large series of each species.” TEHITD LIZARDS OF THE GENUS CNEMIDOPHORUS 3 Strecker (1928), after much field experience, has recently ex- pressed the view that “all of the lizards of the genus Cnemidophorus are exceedingly variable and sometimes the same species differs greatly in color according to the locality in which it is found.” During the progress of this study it has become apparent that the problem of bringing order into the existing taxonomic confusion of Unemidophorus is not as hopeless as many have supposed. A surpris- ingly large amount of material has accumulated in museums—much of this the result of recent collecting. Therefore, very large series of most of the species and subspecies have been available. The situa- tion met has been so strikingly similar to that expressed by Prot. Asa Gray (1863) in reviewing the revision of the oaks (genus Quercus) by the famous botanist, De Candolle, that one must regard his words as being fully as applicable to animals as to plants. It was found that “Where specimens were few, as from countries little explored, the work was easy, but the conclusions of small value. The fewer the materials, the smaller the likelihood of forms interme- diate between any two * * * Where, however, specimens abound * * * here alone were data fit to draw conclusions from.” It is believed that the present study brings the understanding of the genus Cnemidophorus up to date. However, as new series of specimens become available, and additional geographical localities come to be represented, modifications of the ideas of relationships here expressed, and readjustments of the ranges, are inevitable. The writer has been forced to link many forms as sub-species, and, as must have been expected, a reduction in the number of forms recog- nized, rather than an increase, has been necessary. In fact no new species have been found. It is true that nothing can be gained by the “lumping” of doubtful forms, but, on the other hand, much confusion results from the retention of indefinable species and subspecies in the hope that future collections will prove them to be distinct. Certain differences in this genus, such as melanism or the type of mesoptychial scutellation, seem to be largely environmental in nature, and this kind of character subjects a widespread species to seemingly endless variation, as well as to confusing geographic repetition. The writer has prepared this work in conformity to the general principle stated by Blanchard (1921), namely, that “ap- parent constancy throughout a definite geographic range is. suffi- client to warrant the recognition of a name.” Intergradation, as evidenced by a significant degree of morphological approach or over- jap, when confined to a point or line of geographical contact between the two or more diverging populations concerned, has been the criterion for the recognition of subspecies. However, some modi- fication of this idea must be used in the treatment of insular races 4 BULLETIN 154, UNITED STATES NATIONAL MUSEUM that are based only on color or color pattern. A number of island subspecies are recognized here, but none of these vary greatly from their mainland prototypes in scutellation or in bodily proportions. In these cases an apparently constant difference in coloration is usu- ally maintained by each form. Thus, two or more geographical races, which are obviously closely related, may show a really funda- mental color divergence or evolution in spite of a continued uniform- ity in other physical characteristics. The amount of evolution undergone is to be judged largely by quantitative methods. If the color or pattern differences are great and all color stages in the life history of each supposed form are strikingly distinct, the divergence is held to have full specific value, but if these differences are small and show no great break from the parent stock, it is evident that a full specific designation for them would hide the true state of their evolutionary divergence. Popu- lations belonging to the latter type are in reality “mere color varieties,” as they have frequently been styled, and in order that their true value may not be overemphasized and their relationships thus obscured, they have been given subspecific rank in the following pages even though their free interbreeding be now prevented by one or more water barriers. All of the island subspecies recognized in this paper have apparently formed a separate population for only a short time geologically, and the full species have apparently formed an isolated colony for a much greater length of time. If the iso- lation of these subspecies continues, many of them will presumably become full species with the passing of another geological epoch, or less. To give them full specific rank now, however, would obviously be a purely theoretical anticipation, rather than a logical interpre- tation, of nature as it exists to-day. At times overlapping color phases between insular forms appear on intermediate islands. In such cases it is usually necessary to abandon the name for the central group, retaining the two sufficiently differentiated colonies on the neighboring islands as subspecies. The placing of the synonym under the name of the nearest geographical form has almost always been followed, although the actual basis for this act is found, of course, only by a careful comparison of the characteristics of the biological units concerned. In Cnemidophorus, an admittedly plastic genus, it is apparent that colonies of the same general stock are much more homogeneous on small islands than on larger ones, and that insular forms as a whole show a smaller amplitude of variation than do mainland types. The explanation of this may be reached, perhaps, by a brief consideration of the environment. From this study it seems logical to assume that a modern wide ranging species has met with and tolerated a large variety of habitat TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS o conditions. Furthermore, it has been forced to adjust itself to en- vironmental modifications brought about by the extension of its range through its own migrations or by the agency of geological change. Throughout the area of its distribution the individuals concerned must have reached a satisfactory physiological equilibrium with the environment—at least in regard to such prime factors as temperature, moisture, food, shelter, and breeding sites. I ailure to do this would have meant restriction of the range, followed by a steady decrease in the number of individuals, and ultimately it would have spelled extinction. Environmental and mutational differences are usually shght in geographically adjacent populations which show a continuous distribution. The appearance of these differences gives inception to “local races.” Aithough such units are generally not worthy of even subspecific recognition, it appears that they may possess a high degree of evolutionary significance. Thus, it seems logical to beheve that the complete isolation of one of these local phases is often the most important step in that benign process which culminates in the birth of a new taxonomic entity. In fact it ap- pears from this study that at least the greatest percentage of new taxonomic forms do arise only after continued isolation from the parent stock, during which time even slight mutations may have be- come of real significance through inbreeding and accretion. Because of the fact that island forms are more homogenous it seems permissible to diagnose certain new insular species, or to retain those already described, on a smaller sum total of differences than that necessary to discriminate between mainland populations of a similar stock. As previously mentioned, the one criterion to be used in mak- ing a separation is constancy. When the discrimination is between a mainland form and a recently isolated island type, however, it is often extremely difficult to find reliable characters upon which to base a separation. Here the island population may, even after a very careful comparison with representatives from the neighboring mainland, present a seemingly distinctive set of characters. This may cause one not thoroughly familiar with the actual range of variation of the mainland stock to lescribe a new insular species (or subspecies) whose representatives may prove to be nothing but duplicates of those of the original form which dwell in more or less distant regions where there are, perhaps, similar bionomic conditions. In the present instance specimens of the commoner forms have been personally examined at all of the leading American museums both east and west and specimens of the rarer forms and the more critical material have been freely loaned for examination at a central point. This method has given the advantage of comparison of pertinent specimens without the disadvantage on the part of museums at least, 6 BULLETIN 154, UNITED STATES NATIONAL MUSEUM of packing, invoicing, and shipping enormous lots, a procedure that is well nigh impossible in many cases. The assembling of data upon many large series not previously studied, and upon many more not previously correlated, has helped greately in working out the distri- bution and generic relationships of the various forms. A total of 12,676 lizards have been examined during the progress of this study, including 59 of the 86 types here assigned to the genus. Forty of the 59 types in the American museums are in the collection of the United States National Museum at Washington, and the rest are scattered, but in the leading museums of the country. Faunal lists have been consulted extensively for records, and the reports of each species, 1f within the known range, have been ac- cepted. In the following pages the detailed locality records are given for each species. If the report is based on a publication, the latter is cited just after the name of the locality. If specimens sup- porting the record have been examined by the writer, the name of the museum or museums holding them have been added in abbrevi- ated form (see p. 8 for chart of abbreviations). Thus, at a glance one may learn the bas/s for each report presented. Tt is a pleasure for the writer to avail himself of this opportunity to extend his grateful acknowledgments to those institutions and individuals who have cooperated with him in this study. He is par- ticularly indebted to Dr. Frank N. Blanchard, under whose general direction the work has been prepared, for helpful suggestions and criticisms tendered during the preparation of the manuscript. This bulletin, it may be added, forms one of the series of contributions from the Zoological Laboratory of the University of Michigan. The authorities of all of the museums visited during the progress of this enterprise have made their collections available for study and have placed every facility at the writer’s disposal. In this con- nection his appreciation is extended to Dr, Leonhard Stejneger and Miss Doris M. Cochran of the United States National Museum; to Dr. Barton W. Evermann and Mr. Joseph R. Slevin of the California Academy of Sciences; to Dr. Thomas Barbour and Mr. Arthur Loveridge of the Museum of Comparative Zodlogy; to Dr. G. K. Noble and Mr. Clifford H. Pope of the American Museum of Natural History; to Dr. Alexander G. Ruthven and Mrs. Helen T. Gaige of the Museum of Zodlogy of the University of Michigan; to Mr. Karl P. Schmidt of the Field Museum of Natural History; to Mr. H. W. Fowler of the Academy of Natural Sciences of Philadelphia; to Dr: Joseph Grinnell and Dr. Jean Lindsdale of the Museum of Verte- brate Zoology of the University of California; to Dr. J. O. Snyder and Mr. G. S. Myers of the Stanford cen Museum; to Mr. M. Graham Netting of the Carnegie Museum; to Mr. J. K. Strecker of the Baylor University Museum; to Mr. L. M. Klauber of the TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 1 San Diego Society of Natural History; to Mr. C. D. Bunker of the Kansas University Museum; to Dr. Minna E. Jewell and Mr. How- ard K. Gloyd of the Kansas State Agricultural College; and to Dr. Edward H. Taylor of Kansas University for the permission to ex- amine a substantial collection of unreported specimens from the Western States. Yspecially fine loans have been made by the following institu- tions: The United States National Museum (over 1,000 specimens), the American Museum of Natural History, the Field Museum of Natural History, the Museum of Comparative Zoology, the Cali- fornia Academy of Sciences, the Carnegie Museum, and_ the Museum of Vertebrate Zoology of the University of California. Additional loans have been made by Dr. A. I. Ortenburger of the Oklahoma University Museum, Dr. Vasco M. Tanner of Brigham Young University, Dr. J. E. Guthrie of Iowa State College, and Dr. W. L. Burnett of the Colorado State Agricultural College. The writer wishes to express here his appreciation to his wife, May Danheim Burt, who has accompanied him on all of his visits to the museums mentioned above, and has rendered invaluable aid in recording and computing data from the specimens examined. Her interest in the work has been scarcely less than that of the writer himself, and he does not hesitate to say that he feels that her opinions and observations (which have at times differed from his own) have had a great stabilizing influence on the enterprise as a whole. In connection with the review and interpretation of the literature, valuable aid has been given by various persons and agencies. To these the writer is extremely grateful. Especial acknowledgement is here extended to Dr. Peter O. Okkelberg for the translation of the important Danish articles written by Reinhardt and Liitken (1861, 1862), and to Dr. Carl L. Hubbs for checking the agreement of the specific and subspecific names used in this work. Mr. H. W. Parker has kindly furnished a list of the type specimens of Cnemidophorus in the collection of the British Museum as well as specific information concerning the type of Cnemidophorus vittatus Boulenger. Data pertaining to the type of Cnemidophorus roeschmanni Werner have been obtained through the kindness of Herr P. de Grijs of the Hamburg Museum. The author is deeply indebted to Miss Doris M. Cochran of the United States National Museum for the illustrations of color pat- terns, to Mr. M. Graham Netting of the Carnegie Museum for taking the photographs from which the line drawings were made, and to May Danheim Burt for the preparation of the distributional maps and line drawings. Ss BULLETIN 154, UNITED STATES NATIONAL MUSEUM In the designation of the types of the various forms and in con- nection with the distributorial records, the names of collections have been abbreviated as follows: A. NV. S. P., Academy of Natural Sciences of Philadelphia; A. 17. V. H., American Museum of Natural History; Blanchard, private collection of Frank N. Blanchard; B. Y. U., Brigham Young University, Provo Utah; 6:77, Wa British Museum of Natural History; Burt, collected by the writer— all to be transferred to one of the leading American museums; Bay- lor, Baylor University, Waco, Texas; Carnegie, Carnegie Museum, Pittsburgh, Pa.; Colo. Agrt. Coll., Colorado State Agricultural Col- Jege, Fort Collins, Colo.; 7. M. N. H., Field Museum of Natural History; G/oyd, private collection of Howard K. Gloyd; Jowa State, Towa State College, Ames, Iowa; Jewell, private collection of Minna K. Jewell; A. S. A. C., Kansas State Agricultural College, Man- hattan, Kan.; A. U., Kansas University Museum, Lawrence, Kan.; Klauber, private collection of L. M. Klauber; Los Angeles Mus., Los Angeles Museum; d/ich., Museum of Zodlogy, University of Michigan; J. C. Z., Museum of Comparative Zodlogy; WM. V. Z. U. C’., Museum of Vertebrate Zoology, University of California; Ox/a., Oklahoma University, Norman, Okla.; Ottawa Univ., Ottawa Uni- versity, Ottawa, Kan.; San Diego Soc. Nat. Hist., San Diego Society of Natural History, San Diego, Calif.; Stanford, Stanford Univer- sity; Zaylor, private collection of Edward H. Taylor:; U. S. NV. M., United States National Museum; Washburn, Washburn University, Topeka, Kan. References in the text are to author, year and page, and are to be found in full in the bibliography. Many references appear in the synonymies that are not included in the bibliography, however. HISTORICAL REVIEW Certain lizards of the genus Cnemidophorus have been known to naturalists since the early part of the eighteenth century. Perhaps the first to be carried from the New World were /emniscatus and muitnus. These interesting forms were figured by Seba in his his- toric work of 1734 and 1735. The first appeared as Lacerta lemnis- cata in the tenth edition of the Systemae Naturae, but the second was overlooked until 1768 when it was named by Laurenti. Only one other species of Cnemidophorus was made known by Linnaeus, namely, the Lacerta sexlineata of 1766. During the period between 1766 and 1839 the generic position of the two Linnaean species of Cnemidophorus was much varied. Although often referred to the large and unwieldly genus Lacerta, these early forms were also placed at times in the Seps of Laurenti, the Ameiva of Meyer, and 1To be presented to the Kansas University Museum, TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 9 the Ze‘us of Merrem. OCELLIFER---------- isz MURINUSSeee=eeee oe e | ARUBENS IS---~------ 8 7 \ \ oy \ \ es | > | FIGURE 11.—MAP SHOWING DISTRIBUTION OF FORMS GF LEMNISCATUS GROUP ships is given under the affinities of minus (p. 51) and in the general discussion at the end of this work (pp. 251-260). Arubensis of Aruba Island is intermediate, both in geographical position and in characters, between the insular murinus and the mainland lemniscatus. Its brachial scutellation is much coarser than that of typical murinus but much finer than that of typical lemnis- TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 5 catus, and its longitudinal rows of ventral plates are reduced to eight, from the 10 to 12 ventralled condition of mwrinus, as are those of all of the remaining species of Cnemidophorus. The range in the number of femoral pores for murinus has been found to be from 27 to 45, for arubensis 27 to 33, and for lemniscatus from 15 to 29; thus, again, avwbensis is transitional. The pattern of murinus con- sists essentially of a uniform dorsal ground color, broken only by large, round, white spots on the sides, although a few young speci- mens from Curacao, which is geographically adjacent to Aruba, show the suppression of lateral spots and the assumption of more or less distinct dorsal stripes as do juveniles of arwhensis. This process proceeds so far in arubensis that the adult females are often found with stripes and from these to the perfectly striped young of /emnis- catus is but a step. The latter form often reverts to the spotted, unlined, condition in the adult male, developing enlarged blue spots of the sides and at times approaching adult arwbensis in general appearance. Also, the largest adults of /emniscatus are smaller than adults of arubensis and murinus, and the additional derivatives, nigricolor and ocellifer, are even smaller than specimens of lemniscatus. Lemmiscatus is central in relation to the other forms of its group and, as a wide-ranging, plastic entity, has had ample opportunity to give rise to other forms. It is found from Guatemala in the north to northern Brazil in the south. In Brazil its stock has evidently given rise to ocellifer, which has been rendered very distinct by a marked reduction in the number of femoral pores. These openings have changed from a minimum of 15 in lemniscatus to a maximum of 12 in ocellifer. The supraocular granules are also modified in ocellifer, extending much farther forward than in lemniscatus. In the island arc just north of Venezuela, extending from Aves Island just east of Bonaire to Margarita Island, nigricolor has pre- sumably risen from Jemniscatus, possibly being derived after isola- tion by partial submergence of the general area. Subsequent sub- mergence has then, perhaps, isolated the modern island populations of nigricolor from each other. The transition from /emniscatus to nigricolor is still shown by intermediate specimens on Margarita Island which tend to lose all stripes at a very early age and to have the rounded spots on the sides reduced in size. With the exception of an average diminution in the keeling of the subcaudal scales in nigricolor, the form is structurally practically identical with lemniscatus. It may be said here that the break in relationships shown between the lizards on Bonaire (murinus) and Aves Islands (niqricolor), whose habitats are geographically adjacent, may be due to an ancient rift between these two points, 56 BULLETIN 154, UNITED STATES NATIONAL MUSEUM The preservation of the primitive types, murinus and arubensis, on islands, instead of on the mainland, is not without precedent. It may be explained by the theory that the ancestral murimus stock was once a mainland type, but that with topographical changes in the region north of Venezuela, this stock was isolated on the islands, Bonaire, Curacao, and Aruba, while on the mainland is was changed to the progressive ancestral /emniscatus stock. Since Bonaire is farthest from the mainland, it was probably isolated first, and it is not surprising that the presumably least specialized modern group of individuals is to be found here. Before the isolation of Curacao and Aruba, the ancestral murinus stock apparently became modified nigricolor lemniscatus Common stock of nigricolor and lemniscatus ae Ancestral lemniscatus stock Ancestral murinus stock ocellifer modified toward lemniscatus ; ; ae murinus (On Curacao) murinus (On Bonzire) Ancestral mMurinus stock FIGURE 12.—DIAGRAM OF THE SUPPOSED RELATIONSHIPS WITHIN THE LEMNISCATUS GROUP toward lemniscatus and this, if true, will account for the present transitional nature of these particular insular populations. The writer’s interpretations of the genetic relationships within the lemniscatus group are expressed by the foregoing diagram. THE DEPPII GROUP CNEMIDOPHORUS DEPPII DEPPII (Wiegmann) MANY-LINED RACE-RUNNER 1834. Cnemidophorus deppii WieEGMANN, Herpetologia Mexicanna, p. 29, (type locality, ‘“‘ Mexico”’).—Gray, Cat. Liz. British Mus., 1845, p. 22.—Bocovrt, Miss. Sci. Mex. et Amer. Cent., vol. 3, 1874, p. 281, pl. 20ce, figs. 5, 5a-d; pl. 20d, figs. 1, 1a-b—O’SHAuGHNEsSSy, Zool. Rec. for 1878, vol. 15, 1880, “ Reptilia ”, p. 7—Sumicurast, La Naturaleza, ser. 1, vol. 6, 1884, p. 39.— GARMAN, Bull. Essex Inst,, vol. 16, 1884, p. 138.—BouLEeNcER Cat. Liz. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 57 British Mus., vol. 2, 1885, p. 371-—-GUNntTHER, Reptilia and Batr., in Biol. — Centr.-Amer., 1885, p. 27.—Corr, Bull. U. 8. Nat. Mus., vol. 32, 1887, p. 44.— ATKINSON, Ohio Nat., vol. 7, 1807, p. 155—RutTuven, Zool. Jahrb., vol. 32, 1912, p. 322.—Scumipt, Publ. Field Mus. Nat. Hist., Zool. Ser., vol. 12, 1928, p. 198.—MertTEeNS, Abhandl. u. Ber. Mus. f. Naturw. Ver. Magdeburg, vol. 6. 1930, p. 158. 1860. Cnemidophorus decemlineatus HAattoweE nt, Proc. Acad. Nat. Sci. Phila., p. 482, (type locality, “ Nicaragua”; 12 co-types, U. S. Nat. Mus. No. 6058; C. Wright, collector) —BotLencer, Cat. Liz. British Mus., vol. 2, 1885, p. 360. 1863. Cnemidophorus deppei Corr, Proc. Acad. Nat. Sci. Phila., p. 104.—Perrers, Monatsb. d. Konigl. Akad. Wiss. Berlin, 1869, p. 63.—GUnrHeEr, Zool. Ree. for 1869, vol. 6, 1870, p. 111.—Sumicurast, Bull. Soc. zool. de France, vol. 5d, 1880, p. £77.—-F1scHer, Oster-Progr. Akad. Gymn. Hamburg, 1883, p. 2.— Corr, Ann. Rept. U. 8. Nat. Mus. for 1898, 1900, p. 568.—Gapow, Proc. Roy. Soe. London, vol. 72, 1903, p. 113; Science, vol. 22, 1905, p. 688; Proc. Zool. Soe. London, 1905, p. 195; Proce. Zool. Soc. London, 1906, p. 309; Through Southern Mexico, Witherby and Co., 1908, pp. 1389, 166, 181, 2384; Zool. Jahrb., vol. 29, 1910, p. 706.—RutTHVEN, Rept. Mich. Acad. Sci., vol. 14, 1912, Dee2ols 1877. Cnemidophorus lativitiis Copr, Proc. Amer. Philos. Soc., vol. 17, p. 94, (type locality, ** Tuchitan, Tehuantepec,’ Oaxaca, Mexico; type specimen, U.S.N.M. No. 30227; F. Sumichrast, collector) ; Proc. Amer. Philos. Soc., vol. 18, 1879, p. 270.—O’SHAUGHNESSY, Zool. Rec. for 1877, vol. 14, 1879, “ Reptilia,” p. 6—SumicHrast, Bull. Soc. zcol. de France, vol. 5, 1880, pp. 1635 U0 1877. Cnemidophorus lineatissimus Corr, Proc. Amer. Philos. Soc., vol. 17, p. 94 (type locality, ‘‘ Colima and Guadalaxara,’ Mexico; 20 cotypes, U.S.N.M. Nos. 24937-24940, 32299-32314; Mr. Hoge, collector).—O’SHAUGHNESSY, Zool. Ree. for 1877, vol. 14, 1879, ‘* Reptilia,” p. 6—Gwtnruer, Reptilia and Batr., in “ Biol. Cent.-Amer.,” 1885, p. 27. 1892. Cnemidophorus deppei deppei Corr, Trans. Amer. Philos. Soc., vol. 17, p. 31—Burt, Proc. Biol. Soc. Wash., vol. 42, 1929, p. 154. 1892. Cnemidophorus deppei decemlineatus Corr, Trans. Amer. Philos. Soc., VOL ps al. 1892. Cnemidophorius deppei lineatissimus Corr, Trans. Amer. Philos. Soc., vol. 17, p. 31; Amer. Nat., vol. 30, 1896, p. 1024; Ann. Rept. U. S. Nat. Mus. for 1898, 1900, p. 1232. 1894. Cnemidophorus alfaronis Corr, Proc. Acad. Nat. Sci. Phila., p. 199 (type locality, ‘‘“San Mateo, Costa Rica”; type specimen, A.M.N.H. No. 163815; A. Alfaro, collector).—BouLencEr, Zool. Rec. for 1894, vol. 31, 1895, * Reptilia,” p. 23. 1897. Cnemidophorus deppii lineatissimus VAN DENBURGH, Occas. Pap. Calif. Acad. Sci., vol. 5, p. 463. 1910. Cnemidophorus immutabilis WERNER, Mitt. Nat. Hist. Mus. Hamburg, VOL ates Like 1928. Cnemidophorus deppi ORTENBURGER, Mem. Un‘y. Michigan Museums, vol. 1, p. 140. Systematic notes—The many-lined race runner was described by Wiegmann (1834!°) from an 8-striped Mexican specimen. From 10 Since the original description is not generally available, it is thought worth while to quote Wiegmann’s Latin diagnosis in full: “‘Squamis menti parvis, gulae intermediis majoribus ; sq. mesoptychii mediocribus, imbricatis, rhombeis ; seuto labii Ssuperioris primo denticulato ; superne cineraceus, vittis dorsi octo pallide viridibus, ad latera immaculatus.” 2306—31 D 5S BULLETIN 154, UNITED STATES NATIONAL MUSEUM the description it is obvious that the type represents the present form, and it seems important that it was recognized that there is a very close resemblance between deppii and lemniscatus, the chief difference being the normal possession of three large parietals and three supraoculars in deppii as opposed to the normal possession of five large parietals and four supraoculars in /emniscatus. ‘The color is in many cases practically identical. Cnemidophorus decemlineatus was described by Hallowell in 1860 from a series of 10-lined specimens from Nicaragua. Since 10-lined specimens are common throughout the range of deppiz and occur very frequently in Colima and Nayarit in the northern part of it, the suppression of decemlineatus seems entirely justified. A specimen from Oaxaca was described as lativittis by Cope in 1877. The type has eight lines and this resemblance to octolimeatus (= perplexus) caused Cope to cite differences from that form in his diagnosis. Since these differences were all toward deppii which is widespread in the coastal region of Oaxaca, lativittis was early reduced to its synonymy. The specimens from Colima and Jalisco, as well as those examined by the writer from Nayarit, usually differ from specimens from Oaxaca and Vera Cruz by having ten or eleven pale lines on the back as do specimens from Nicaragua. This variation caused Cope (1877) to describe a series of such northern variants as /ineatissimus. Since 10 or 11 lined deppii are sometimes found in Oaxaca and Vera Cruz and since they also occur normally in certain parts of Nicaragua at least, there is nothing geographically distinctive in lineatissimus. ‘Therefore, it, too, has been relegated to the synonymy of deppii. Some abnormal specimens of deppii from San Mateo, Costa Rica, were described by Cope in 1894 as alfaronis. These were typical depp in coloration, having 9 dorsal lines, but differed in scutella- tion by the presence of 4 supraoculars instead of 3. Since this anom- aly occurs now and then in deppii everywhere, as remarked by Gadow (1906, 309) and indicated below (see p. 60), alfaronis can not be retained as a distinct entity. DPiagnosis—The many-lined race runner is distinguished by its small maximum size (usually less than 85 mm. from snout to anus) and the usual presence of 3 supraoculars, 3 parietals, granular post- antebrachials and weak caudial keels. Specimens are seldom, if ever, unicolor, and may be either spotted or lined. The normal young possess from 6 to 13 well-defined, white stripes, which are usually straight, but may be more or less wavy. With development spots may appear in the lateral interspaces, but they never spread to form cross-bars as in gulards. The lines on the sides, especially the lower TERIID LIZARDS OF THE GENUS CNEMIDOPHORUS 59 pair, may disappear with development, or the 3 dorsal stripes may united into a single broad, middorsal streak, and at times all of the stripes are very faint, almost obsolete. Although they may usually be diagnosed at once, examples of dep- pii ave sometimes so close to guttatus that separation is almost im- possible. There are apparently no differences in scutellation. How- ever, most individuals of guttatus exceed the maximum length of deppii indicated above, and although there are usually two distinct, lateral lines at the base of the tail in depp7/, there is usually only one in guttatus. Also, the femora of deppii are usually covered by distinct, rounded, light spots, while those of guftatus are usually unicolor or obscurely marked. Anal spurs, although often found in the adults of /emniscatus, are universally absent in deppii. Description—Snout moderately elongate; nostril anterior to nasal suture; anterior nasal usually not in contact with second upper labial; supraoculars normally 3; superaocular granules usually small, forward extent variable; frontoparietals normally 2; parietals 3; anterior gulars moderate to large, graded, largest centrally ; posterior eulars smaller, contrasted with anterior gulars; mesophychial scales moderate to large, largest centrally, or medio-laterally, graded and smaller laterally, in 2-5 rows; postmesoptychial granules minute, sometimes extending to edge of posterior mesoptychial fold. Body elongate; ventral plates in 8 longitudinal and 380-38 trans- verse rows; dorsal granules minute to moderate; limbs well devel- oped; brachials 3-8; antebrachials 2-4; brachials and antebrachials more or less continuous at a point of contact; postantebrachials small or slightly enlarged; femorals 4-9; tibials 2-4; femoral pores 12-23; tail elongate, tapering; subcaudals smooth; lateral and dorsal ‘audals weakly keeled, shghtly oblique and moderate in size, keels usually irregularly arranged. Coloration variable; lower surfaces white to blue-black, often slaty or slaty-blue, especially in the young; lower surfaces of tail usually white or yellowish, upper surface brown, blue, gray or olivaceous, usually not spotted or reticulated; tail usually with two narrow, diminishing, lateral stripes on each side of its base; femora usually strongly spotted or reticulated with white, especially in the young, but sometimes nearly unicolor; back usually lined, rarely spotted; sides spotted, lined and spotted, or only lined; if fully lined, 6 to 13 stripes present; width of stripes variable, stripes usually straight, seldom wavy, dorsal pair often converging on the median line posteriorly and anteriorly to produce an elongate, double- pointed oval, and sometimes united with the middorsal stripe to produce a broad, light streak; dorsal ground color usually blackish or gray, but sometimes brownish. 60 BULLETIN 154, UNITED STATES NATIONAL MUSEUM A series of 1023 specimens of depp has been examined. Data on a representative series of 700 examples are as follows: Body 24-86 mm.; tail, 54-207; total length, 86-288; width of head, 4.0-12.5; length of hind leg, 20-55; supraoculars 3 in 612 speci- mens, 3-4 in 40 specimens and 4 in 48 specimens; supraocular granules extending forward to the anterior border of the fourth supraocular in 1 specimen, to the middle of the third supraocular in 601 specimens, to the anterior border of the third supraocular in 46 specimens, to the middle of the second supraocular in 9 specimens and to the anterior border of the second supraocular in 2 specimens. Variation —Gadow (1903, p. 113) called attention to the fact that not over 6 per cent of deppii have less than 9 longitudinal hght lines on the body. The occurrence of a 6-lined specimen is very ‘are, but 8-lined ones are found more frequently. Gadow (1906, p- 295) wrote that “the greatest number of stripes occur in old specimens, and this fact is not due to the others having been weeded out, since many-striped young are not relatively, but positively, rare.” The writer finds that the young and the adults usually show the same number of stripes and sees no reason why more should be found in adults. However, it is just possible that Gadow confused young, six-lined specimens of gudaris with those of deppii. Abnormalities in the striping frequently occur in Nayarit, Jalisco and Colima for here the usual dorsal stripes, although often dis- tinct, frequently combine to produce a single, widened, heavy, con- spicuous middorsal line. This aberration is seldom seen elsewhere. As maintained elsewhere (p. 58), regional variation in the number of stripes is common, but has little taxonomic significance because of its repetitive character. Gadow (1906, p. 309) listed the supraoculars of 152 specimens as normally three, but said that there were 10.0 percent of exceptions. This is close to the writer’s figure of 9.7 percent, which is based on 700 examples. Range.—The many-lined race runner occurs in the lower levels along the Mexican coast southward from Nayarit and Vera Cruz, and is found in Central America as far south as Costa Rica. It probably does not extend inland above an elevation of 4,000 feet. The report of deppii from Caracas, Venezuela, given by Boul- enger (1885), and copied by others, was probably based on a young specimen of lemniscatus. The available records are presented below in an alphabetical series by countries, and under them by the largest political sub- divisions. COSTA RICA.—Prov. of Alajuela (San Mateo, Cope, 1894, p. 199, A.M.N.H.). Comarca de Puntarenas (T7irwes, U.S.N.M.). TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 61 GUATEMALA.—Dept. of Guatemala (Guatemala, U.S.N.M.; Hacienda Neja, M.C.Z.). Izabal (Los Amates, Atkinson, 1907, p. 155). Peten (General Report, U.S.N.M.). Zacapa (Gualan, At- kinson, 1907, p. 155, U.S.N.M., A.N.S.P.; Zacapa, U.S.N.M.). HONDURAS.—Dept of Cortes (San Pedro, US.N.M.; West of San Pedro, ¥.M.N.H.). Tegucigalpa (Hacienda Santa Ana, ¥’.M.N.H.). Valle (Road above Pespore, ¥.M.N.H.). 100 90 FIGURE 13. MAP SHOWING LOCALITY RECORDS FOR CNEMIDOPHORUS DEPPIIL DEPPII (REPORTS BY LARGEST POLITICAL SUBDIVISIONS) MEXICO.-—States of Chiapas (Zonala, U.S.N.M.). Colima (Campos, Los Angeles Mus.; Colima, Cope, 1877, p. 94, U.S.N.M.. ASMUN.H., M.C.Z., Mich.; Hast of Colima, AM.N.H; South of Colima, A.M.N.H.; La Quinta, A.M.N.H.; Los Tapestes, A.M.N.H.; Manzanillo, Los Angeles Mus.; San Cayetana, A.M.N.H.; 6 miles east of Tecoman, A.M.N.H.). Guerrero (Acapulco, Los Angeles Mus. ; Ayutla, Gadow, 1906, p. 277; Balsas, Gadow, 1906, p. 277, F.M.N.H.; Cacahuamilpa, M.V.Z.U.C.; Chilpancingo, Gadow, 1906, p. 277; Cocoyul, Gadow, 1905, p. 195; Los Cajones, Gadow, 1906, p. 277; Tierra Colorado, Gadow, 1906, p. 277; T/apa, U.S.N.M.; Totolapan, Gadow, 62 BULLETIN 154, UNITED STATES NATIONAL MUSEUM 1905, p. 195). Jalisco (La Tres Marietas Islands, middle island, C.A.S.; Vallarta, A.M.N.H.; Valley near Guadalajara, Cope, 1877. p. 94, U.S.N.M.). Michoacan (Alvarado, M.C.Z.; La Salada, U.S.N.M.; Mescala, Gadow, 1906, p. 277). Nayarit (Minamon, U.S.N.M.; San Blas, Van Denburgh, 1897b, p. 463, U.S.N.M., A.M.N.H.). Oaxaca (Chivela, A.M.N.H., M.C.Z.; Juchitan, Cope, 1877, p. 95, U.S.N.M.; Miahuatlan to Ejutla, A.M.N.H.; Mixtequillo, C.A.S., A.M.N.H.; Oaxaca, M.V.Z.U.C.; Salina Cruz, Gadow, 1906, p. 277, A.M.N.H.; San Carlos, Gadow, 1906, p. 277; San Domingo, Gadow, 1906, p. 277; San Geronimo, A.M.N.H., F.M.N.H.; San Luis Allende, Gadow, 1906, p. 277; Santiago Astata, A.M.N.H.; Tapana- tepec, M.C.Z.; Tehuantepec, Gadow, 1906, p. 277, U.S.N.M.; 3 males north of Tehuantepec, A.M.N.H.; West Tehuantepec, Cope, 1887, p. 44; Tequesiatlan, Gadow, 1906, p. 277). Vera Cruz (Agua Fria, Gadow, 1906, p. 277; Chichicaxtle, U.S.N.M.; Coatzacoalcos Bay, U.S.N.M.; Cwatotlapan, Mich.; El Potrero, M.C.Z.; Mandinga, A.M.N.H.; Otopa, Ruthven, 19126, p. 231, F.M.N.H.; South of Panteon Viego, A.M.N.H.; Perez, Ruthven, 19126, p. 231, F.M.N.H..; Sand Dunes 2 miles south of Vera Cruz, A.M.N.H.; San Francisco, Ruthven, 19120, p. 231; San Juan Hvangelista, Gadow, 1906, p. 277; Vera Cruz, Ruthven, 19120, p. 231, A.M.N.H., C.A.S., F.M.N.H.). NICARAGUA.—Dept. of Chontales (San Lorenzo, ¥.M.N.H.). Chinandega (Chinandega, M.C.Z.; Corinto, Werner, 1910, p. 27). Granada (Granada, ¥.M.N.H.). Leon (d/omotombo, ¥.M.N.H.). SALVADOR.— (General Report, Gunther, 1885, p. 27). Habitat and habits —Very little is known concerning the habitat of this form and evidently nothing has been published about its habits. According to Gadow (1910, p. 706) deppii reaches a max- imum altitudinal distribution of 4,000 feet in Mexico. The same author (1908, p. 139) found it on the “ open savannahs of Juanita,” and recalled (p. 234) having seen many examples “in the sandy bed of the Tehuantepec River of southern Mexico.” Ruthven (1912a, p. 822), working in Vera Cruz, found this subspecies to be very rare, and wrote as follows: “ The three specimens taken were found in a thicket of low bushes on the savannah near the San Juan River. In this thicket, which was about a hundred meters in diameter, there were scores of Ameiva undulata, but these three specimens of C. deppii were the only ones seen. That they were really rare in this habitat and not merely overlooked was shown by the fact that the thicket was worked thoroughly, and the three specimens were taken on different days in almost exactly the same place.” Enemies —Vhe Mexican whip snake, MJasticophis mentovarius, was mentioned by Gadow (1908, p. 166) as preying upon these lizards. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 63 Affinities —The close relationship existing between deppii and lemniscatus has been emphasized above. For reasons to be given in the general discussion at the end of this work, deppii is presumed to have originated from the /emniscatus stock somewhere in Central America, thus giving rise to the group which bears its name. The closest modern relative of depp7i is evidently cozwmelus with which it is structurally identical. Both are of the same general size. C. guttatus, although agreeing in scutellation and not varying greatly in coloration, differs considerably by its larger size, and is consid- ered to be a somewhat more distant relative. The relationships of cozumelus and guttatus with deppii are discussed in detail below under the affinities of those forms. CNEMIDOPHORUS DEPPIL COZUMELUS Gadow BROWN-BACKED RACE-RUNNER 1906. Cnemidophorus deppei cozumela GApow, Proc. Zool. Soe. London, p. 316 (type locality, “ Cozumel Island, east coast of Yucatan” peninsula, Quintana Roo, Mexico; 4 co-types, B.M.N.H., 1886.4.15.17-20). Diagnosis —The brown-backed race-runner may be distinguished as follows: supraoculars normally 3; parietals 3; postantebrachium covered with small polygones or scutes; anal spurs absent; each side with 2-5 narrow, wavy, irregular, light lines; back unicolor, with an unusually wide, yellowish brown, longitudinal band present; ground color of sides dark reddish brown, in abrupt contrast to color of dorsal band: sides and femora never with conspicuous white or blue spots; lower surfaces white or light blue. This subspecies is easily distinguished from other forms of Cnemidophorus by these characters and is approached only by a phase of deppii which sometimes appears in southern Mexico and adjacent areas to the south, from which it may be separated by the cmusually wide dorsal band and the unusually wavy, irregular, stripes on the sides. Description—Snout moderately pointed; nostril anterior to nasal suture; anterior nasal separated from second upper labial; supra- oculars normally 3; supraocular granules usually not extending for- ward past the posterior border of the second supraocular; fronto- parietals normally 2; parietals normally 3; anterior gulars small to moderate laterally, graded, and with an enlarged patch medially ; anterior and posterior gulars usually abruptly differentiated; pos- terior gulars minute; mesoptychials moderate to large, uniform, or a little smaller laterally, arranged in 2-5 enlarged transverse rows: postmesoptychials minute usually hid behind posterior gular fold. Body moderately elongate; ventrals arranged in 8 longitudinal and 31-36 transverse rows; dorsal granules fine; limbs well devel- oped; brachials 6-9; antebrachials 2-4; brachials continuous with 64 BULLETIN 154, UNITED STATES NATIONAL MUSEUM antebrachials at a point of contact; postantebrachium covered with small or slightly enlarged granules; femorals 6-10; tibials 2-4; femoral pores 15-19; tail elongate tapering; caudals moderate, lateran and dorsal keels weak, and more or less irregularly arranged. SA aa a Si er FIGurRE 14.—COMPARISON OF TYPICAL COLOR PATTERN OF CNEMIDO- PHGRUS DEPPI COZUMELUS (LEFT) AND C, DEPPIL DEPPII (RIGHT). NOTE THE WIDER MIDDORSAL BAND AND WAVY LATERAL STRIPES IN COZUMELUS Coloration distinctive; lower surfaces white or suffused with light blue; tail uniformly brownish above, without dark reticulations; femora usually uniform brownish, spots or reticulations, if present, very obscure; femur usually not striped behind; back with and unusually broad, longitudinal band of uniform ght brown; sides with 2-5 stripes each, these wavy, irregular and narrow; ground color of sides dark reddish brown, in sharp contrast to dorsal color. The following data have been secured from three specimens from TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 65 Cozumel Island U.S.N.M. Nos. 47653-47655): Body, 57-75 mm.; tail, 137-141; total length, 194-216; width of head, 7-10; hind leg, 42-52; supraoculars 3-8 in one specimen, and 3—4 (fourth vestigial) in two specimens; supraocular granules extending forward to the middle of the third supraocular in two specimens, and to the anterior border of the third supraocular in one specimen. The frontoparietals of these three specimens are all abnormally divided and in different ways; the result is three scutes in every case. A single specimen from Mujeres Island (U.S.N.M. No. 47652) is typical in coloration and in other features as well. It is ¢4 mm. long and has a regenerated tail; the head width is 7 mm.; and the hind leg is 44 mm. long. Some other features are: supraoculars, 3-3; supraocular granules extending forward to the middle of the third supraocular; and femoral pores 17-18. The frontoparietal is partly united from behind, but there is a squarish lateral scute split off from each external border posteriorly to produce a total of three scutes. This arrangement is often found in the South American teiid, Dicrodon lentiginosus. In the original description Gadow, using an underline, emphasized the fact that “not only the first but also the second and even the third upper labials are denticulated.” This seems to be a very unim- portant distinction, however, since the same condition is found fre- quently in deppii. In all of the specimens examined by the writer the loreal is in contact with the first supraocular, but the amount of contact varies considerably, and like the denticulation of the upper labials, it is not wholly diagnostic. Range.—Specimens of this lizard have been examined from two islands which le off the east coast of Quintana Roo, Peninsula of Yucatan, in southern Mexico. These are: (Cozwmel Island, Gadow, 1906, p. 316, U.S.N.M.; and Afujeres Island, U.S.N.M.). Affinities —This form is structurally identical with deppi, from which it differs only in coloration. Specimens from the mainland (deppii) sometimes possess a more or less widened middorsal yellow- ish streak which resembles that of sev/ineatus. In these the lateral stripes are usually straight, but occasionally they are more or less wavy. The transition from these to cozumelus is but a step. This is accomplished by the widening of the dorsal band and the modifica- tion of the lateral stripes. Thus, cozwmelus may be considered as a direct derivative of the wide-ranging mainland deppii, from which it has probably been differentiated for a comparatively short time. (or a comparison of the typical color patterns of the two lizards see figure 14.) = 66 BULLETIN 154, UNITED STATES NATIONAL MUSEUM CNEMIDOPHORUS GUTTATUS (Wiegmann) MEXICAN RACE-RUNNER 1834. Cnemidophorus guitatus WIEGMANN, Herpetologia Mexicana, p. 29 (type locality, “* Mexico”; type specimen, Berlin Mus. No. 867).—DuMERILL and Breron, Erp. Gén., vol. 5, 1839, p. 134.—Gray, Cat. Liz. British Mus., 1845, p. 22.—HALLOWELL, Reptiles, Pacific R. R. Surv. (Parke Route), vol. 10, pt. 6. sect. 2, no. 2, p. 23.—GUNTHER, Zool. Rec. for 1865, vol. 2, 1865, p. 148; Zool. Ree. for 1866, vol. 3, 1867, p. 123.—PrtTrers, Monatsber. Berlin Akad. Wiss.. 1869, p.. 68.—GUntuHeErR, Zool. Rec. for 1869, vol. 6, 1870, p. 111.—Bocourrt, Miss. Sci. Mexique et Amer. Cent., vol. 3, 1874, p. 285, pl. 20c, figs. 4, 4a—d— Corr, Proc. Amer. Philos. Soc., vol. 17. 1877, p. 94.—FiscHrEr, Oster-Progr. Akad. Gymn. Hamburg, 1883, p. 2-——GARMAN, Bull. Essex Inst., vol. 16, 1884, p. 13.—BouLENGER, Cat. Liz. British Mus., vol. 2, 1885, p. 370.—GUNTHER, Reptilia and Batr., in ‘“ Biol. Centr.-Amer.,”’ 1885 p. 28—BouLencer, Zool. Ree. for 1885, vol. 22, 1886, ‘ Reptilia,’ p. 14.—FrrrAriI-PEREz, Proc. U. S. Nat. Mus., vol. 9, 1886, p. 195.—Coprr, Bull. 32, U. S. Nat. Mus., 1887, p. 44.—Borrtcer, Kat. der Rept.-Samml. im Mus. Senckenb. Naturforsch. Ges. Frankfurt, teil 1, 1893, p. 75—Corn, Amer. Nat., vol. 30, 1896, pp. 1014, 1021-1022, 1025; Ann. Rept. U. S. Nat. Mus. for 1898, 1900, p. 568.—Gapow, Proe. Zool. Soe. London, 1905, pp. 195, 216—AtTKINSOoN, Ohio Nat., vol. 7, 1907, p. 155.:—Gapow. Through Southern Mexico, Witherby and Co., 1908, p. 128.—RuTHVEN, Rept. Mich. Acad. Sci., vol. 14, 1912, p. 231.— SHERBORN, Index Animalium (1801-1850), pt. xi, 1926, p. 2864. 1839. Cnemidophorus searlineatus (part) Dumerrtn and Breron, Erp. Gen., vol. 5, p. 181—WESTPHAL-CASTELNAU, C. R. des trav. de Cong. sci. de France, tenu & Montpellier en déc. 1868, 1870, p. 19. 1851. Cnemidophorus sexrlineatus guttatus DumMeérin and DuMmMEéERIL, Cat. méth. Coll. Reptiles, Paris, 1851, p. 116.—LicHTeNsrern, Nomenclator Berol., 1856, p. 138.—JAn, Indice Sist. dei. Rettili ed Anfibi Mus. Civ. di Milano, 1857, p. 40. 1862. Ameiva guttata Corr, Proc. Acad. Nat. Sci. Phila., p. 63.—PETERS, Monatsbr. Berl. Akad. Wiss., 1871, p. 399. 1875. Amiva guttata Corr, Journ. Acad. Nat. Sci. Phila., ser. 2. vol. 8, 1876, p. 118. 1877. Cnemidophorus immutabilis Corn, Proc. Amer. Philos. Soc., vol. 17, p. 93, (type locality, ‘* West Tehuantepec,” Oaxaca, Mexico; type specimen, U.S.N.M. No. 380141; F. Sumichrast, collector) ; Proc. Amer. Philos. Soc., vol. 18, 1879, p. 270.—O’SHAUGHNESSy, Zool. Rec. for 1877, vol. 14, 1879, ‘‘ Reptilia,” p. 6.—SuMIcHRAST, Bull. Soc. Zool. de France, vol. 5, 1880, pp. 168, 177; La Naturaleza, ser. 1, vol. 6, 1884, p. 89—BouLrenerr, Cat. Liz. British Mus., vol. 2, 1885, p. 370.— Corr, Bull. U. S. Nat. Mus., vol. 32, 1887, p. 44.—BouULENGER, Proce. Zool. Soc. London, 1898, p. 916.—Gapow, Through Southern Mexico, Witherby and Co., 1908, pp. 166, 181; Zool. Jahrb., vol. 29, 1910, p. T06.— ORTENBURGER, Mem. Univ. Michigan Museums, vol. 1, 1928, p. 140. 1877. Cnemidophorus microlepidopus Corr, Proc. Amer. Philos. Soc., vol. 17, p. 93 (type locality, “West Tehuantepec,” Oaxaca, Mexico; type specimen, U.S.N.M. No. 30187; F. Sumichrast, collector) ; Proc. Amer, Philos. Soe., vol. 18, 1879, p. 270.—O’SHAUGHNESSY, Zool. Rec. for 1877, vol. 14, 1879, “ Rep- tilia,” p. 6—Sumicurastr, Bull. Soc. Zool. France, vol. 5, 1880, pp. 163, 177; La Naturaleza, ser. 1, vol. 6, 1884, p. 39. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 67 S77. Cnemidophorus wricolor Cork, Proc. Amer. Philos. Soe., vol. 17, p. 93 (type locality, “ West Tehuantepec,” Oaxaca, Mexico) ; Proc. Amer. Philos. Soe., vol. 18, 1879, p. 270.—O'SHaucHNEssy, Zool. Rec. for 1877, vol. 14, 1879, “ Reptilia,” p. 6—Sumicurast, Bull. Soc. Zool. de France, vol. 5, 1880, p. 177; La Naturaleza, ser. 1, vol. 6, 1884, p. 39. 1892. Cnemidophorus guttatus guttatus Corr, Trans. Amer. Philos. Soc., vol. 17, p. 32—Gapow, Proc. Zool. Soe. London, 1906, pp. 309, 325. 1892. Cnemidophorus guttatus immutabilis Corr, Trans. Amer. Philos. Soc., vol. 17, p. 31—Gapow, Proc. Zool. Soc. London, 1906, pp. 309, 326. 1892. Cnemidophorus guttatus microlepidopus Core, Trans. Amer. Philos. Soc., VOLeiGop os. 1892. Cnemidophorus guttatus unicolor Corr, Trans. Amer. Philos. Soc., vol. 17, Dome 1903. Cnemidophorus guttatus guitata GApow, Proc. Roy. Soc. London, vol. 72, p. 115. 1903. Cnemidophorus guttatus striata Ganow, Proc. Roy. Soc. London, vol. 72, p. 115 (type locality, ‘“ Isthmus of Tehuantepec,’ Oaxaca, Mexico). 1905. Cnemidophorus striatus Gapow, Proc. Zool. Soc. London, 1905, p. 195. Systematic notes—The Mexican race-runner was described by Wiegmann in 1834 from “ Mexico” and, although the original account’? is incomplete, subsequent workers have elaborated it to definitely fit the present form. It is evident that the type is of the spotted, rather than of the lined or unicolor phases of the species. As described by Gadow (1906, p. 373), the young examples of guttatus, as well as some larger, forest inhabiting, individuals, are dull and frequently even unicolor. It seems that with development stripes or spots appear, usually to partly vanish again. In the case of the striped form, the lines often break into spots posteriorly before becoming obsolete, and then perhaps anteriorly as well. There is much individual variation in regard to these pattern changes. In some regions the lizards apparently retain a distinctly spotted or lined pattern throughout life. The dull phase of guttatus was described as unicolor by Cope (1877), who diagnosed it very fully. The same author recognized his mistake later (1887, p. 44) and listed this form in the synonymy of guttatus. The striped phase of guttatus, which apparently reaches its max- imum development in certain parts of Oaxaca, was called * ¢mmuta- bilis” by Cope (1877). Thus, as remarked by Gadow (1906, p. 326), by the irony of fate 7mmutabilis became the proper name of one of the most variable of lizards. The chief character of this form was * the retention of stripes throughout life.” Gadow (1906, p- 287) wrote that “ (. guttatus and immutabilis were considered as good species by Cope and Boulenger, but intermediate forms are 11 Because of its obscurity, it is thought best to present the original Latin diagnosis of guttatus in full, as follows: “‘Squamis gulae intermediis majoribus squamis mesoptychii magnis, subrhombeis, imbricatis; dorso cineraceo, guttis albidis longitudinaliter adsperso ; lateribus superne nigricantibus, immaculatis, vittaque gemina marginatis.” BULLETIN 154, UNITED STATES NATIONAL MUSEUM 68 NGAGS OL LHSIN WOU SadiuLs NOLLISNVUL DNIMOHS ‘ASVHd GUNIT GHL AO YNIdIIS H V4 % ; ; TVWSuo0d GHL NI NOLLVIUVA ‘SOLVLEAD SauoHaodINAN)J— GT ANNI TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 69 found living in districts of intermediate bionomic conditions, so that now at best they are subspecies, if not local races, or, worse still, only pattern varieties.” It is significant that most of Gadow’s inter- mediate specimens came from southeastern Guerrero, instead of from the Isthmus of Tehuantepec. These “intergrades ” are not geographically intermediate between the spotted eastern quttatus, which Gadow confined to Vera Cruz, and the striped ¢mmutabilis from Oaxaca, but are found isolated on the Pacific slope of Oaxaca where only typical 7mmutabilis is theoretically expected. After the examination of a large series of these lizards the writer feels that the two entities under discussion should no longer stand apart. A1- though certain populations do appear to be moderately well differ- entiated, pattern intermediates are found in many places and in addition the appearance of unicolored young in both phases makes it increasingly difficult to separate one from the other. Moreover, it appears that the lined phase sometimes changes into the spotted one. A specimen typical of ¢mmutabilis was described as microlepidopus by Cope (1877) with the statement that “ This form differs from the typical guttatus in color only and may be the young. The median dorsal region is, however, unspotted.” An examination of the type shows it to be an 8-lined specimen, the dorsal stripes unbroken. Just why émmutabilis and microlepidopus should have been designat- ed as different species in the same contribution is not apparent. Evidently through an oversight, Gadow (1903) described striatus from the type locality of tmmutabilis, citing differences from the spotted guttatus. Soon after this (1906, p. 374), he listed the new form in the synonymy of dmmutabilis. Diagnosis —The Mexican race runner is distinguished by its large maximum size and the usual presence of three supraoculars, three parietals, granular postantebrachials and weak caudal keels. Speci- mens may be unicolor, spotted or striped. If spotted, the spots are usually confined to the dorsal region only. The unicolor specimens are mostly young and are generally gray or blackish instead of brownish as in gudaris. ‘The maximum number of stripes is eight. Examples of deppii are sometimes so close to guttatus that separa- tion is almost impossible, although usually specimens may be diagnos- ed at once. There are apparently no differences in scutellation. The largest deppii seldoin exceed a length of 80-85 mm. from snout to anus, but the larger guttatus specimens always exceed this length and the smaller ones are generally unicolor. Although there are usually two distinct, lateral, light lines on each side of the base of the tail in deppii, there is usually only one in guttatus, and whereas the femora of deppii are often covered by distinct, rounded, light spots, those of guttatus are usually obscurely spotted or reticulated 70 BULLETIN 154, UNITED STATES NATIONAL MUSEUM with white. The unicolor phase probably does not occur in deppi. Anal spurs, although often found in lemniscatus, are universally absent in guttatus. Description.—Snout moderately elongate; nostril anterior to nasal suture; anterior nasal usually not in contact with second upper labial; supraoculars normally 3; supraocular granules usually small, forward extent variable; frontoparietals normally 2; parietals 3; anterior gulars moderate to large, graded, largest centrally; posterior gulars smaller, contrasted with anterior gulars; mesoptychial scales moderate to large, largest centrally or medio-laterally, graded and smaller laterally, in 2-5 rows; postmesoptychial granules minute, sometimes extending to edge of posterior gular fold. Body elongate; ventral plates in 8 longitudinal and 30-40 trans- verse rows; dorsal granules minute to moderate; limbs well de- veloped, strong; brachials 4-7; antebrachials 2-8; brachials and ante- brachials continuous or subecontinuous at a point of contact; post- antebrachium covered with small or shghtly enlarged granules; femorales 5-10; tibials 2-4; femoral pores 17-26; tail elongate, taper- ing; subcaudals smooth; lateral and dorsal caudals smooth or weakly keeled, slightly oblique, and moderate in size, keels irregularly arranged. Coloration variable; lower surfaces white to dark blue, often slaty or slaty blue, especially in the young; lower surface of tail usually white or yellowish, upper surface brown, blue, or olivaceous usually not spotted or reticulated; tail with a narrow, diminishing, lateral stripe on each side anteriorly, seldom with two stripes on each side; femora weakly spotted or reticulated with white; back unicolor, spotted or lined; if unicolor, traces of stripes may be present later- ally; if spotted, lines may or may not be present laterally; if lined, spots may or may not be present dorsally or posteriorly, stripes varying from six to eight in normal individuals, width of stripes variable; dorsal ground color usually blackish or gray, seldom brownish, A series of 448 specimens of the Mexican race runner has been examined, and data from 300 of these may be presented as follows: Body, 35-138 mm.; tail, 130-295; total length, 184-425; length of tail as percentage of total length, 60-73.2; width of head, 6-22; width of head as percentage of body length, 12.0-15.7; hind leg, 26-83; length of hind leg as percentage of body length, 58.0-17.4; frontoparietals partly united into a single scute in 3 specimens, dis- tinct and in two scutes in 292 specimens, and abnormally split into three scutes in 5 specimens; supraoculars three in 279 specimens, 3-4 in 10 specimens, and 4 in 11 specimens; supraocular granules extend- ing forward to the middle of the third supraocular in 113 specimens, to the anterior border of the third supraocular in 78 specimens, to TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS ‘FL the middle of the second supraocular in 74 specimens, and to the anterior border of the second supraocular in 34 specimens. Variation —Gadow (1906, p. 320) reported upon 61 specimens of the present species. He found that the * supraoculars were normally three, but that there are 12-14 percent exceptions.” Calculations from the writer’s data on the 300 specimens discussed above, show less than 6 per cent exceptions. 100 90 eC RR A CF CE ESE ERE EE EE FIGURE 16.—Map SHOWING LOCALITY RECORDS (STATE REPORTS) FOR CNEMIDOPHORUS GUTTATUS Gadow (p. 283) examined the dorsal granules of a series of guttatus from Aqua Fria and found that they vary in number from 100 to 180 across the back. Peters (1871, p. 399) wrote that the type specimen has the frontal shield transversely divided behind. This condition is normal in Ameiva bifrontata bifrontata and its allies from northern South America, but is wholly abnormal in guttatus. Aside from the tendency for the adults of the Mexican race-runner to possess a spotted pattern in Eastern Mexico, and to possess a striped pattern in Western Mexico, as discussed in the systematic notes, little geographical variation is evident. (2 BULLETIN 154, UNITED STATES NATIONAL MUSEUM Range.—The Mexican race-runner is apparently confined to the mainland of Mexico and occurs along the Gulf coast of Vera Cruz, throughout the region of the Isthmus of Tehuantepec, and along the Pacific coast from Chiapas to Michoacan. Its detailed distribution follows alphabetically by states: CHIAPAS.—(Tonala, U.S.N.M.; Mowntains near Tonala, Uss. NM) GUERRERO.— (Acapulco, U.S.N.M., A.M.N.H.; Ayutla, Gadow, 1906, p. 809; Chilpancingo, Gadow, 1906, p. 309; Cocoyul, Gadow, 1906, p- 309; a ee Gadow, 1906, p. 309; Tierra Colorado, Gadow, 1906, p. 309; Votolapan, Gadow, 19050, pz 195). mC ee MCyZe,. Miche): OXACA:.—(Chacalpa, A.M.N.H.; Chivela, A.M.N.H., M.C.Z.; Mixtequillo, A.M.N.H., C.A.S.; Mountain, Giengda, A.M.N.H.; Oaxaca, M.V.Z.U.C; Sadani, AM.N.H.; Salina Cruz, Gadow, 1906, p. 309; Salina Cruz Cemetary, A.M.N.H.; West Salina Cruz, A.M.N.H.; San Geronimo, A.M.N.H., F.M.N.H.; San Luis Allende, Gadow, 1906, p. 309; San Mateo del Mar, Gadow, 1906, p. 309; Santiago Astata, A.M.N.H.; Santo Domingo, Boet:ger, 1893, p. 75; Tapanatepec, M.C.Z.; Tehuantepec, U.S.N.M.; 3 miles north of Tehuantepec, A.M.N.H.; West Tehuantepec, Cope, 1887, p. 93; Tequesixtlan, Gadow, 1906, p. 309). VERA CRUZ—(Coatzacoalcos Bay, U.S.N.M.; El Potrero, M.C.Z.; Jalapa, Cope, 1887, p. 44, M.C.Z.; Za Antigua, Ruthven, 19126, p. 231, F.M.N.H.; Orizaba, Cope, 1887, p. 44; Otopa, Ruthven, 19120, 231, F.M.NE: Perez, Ruthven 19120, p. 231, F.M.N.H.; A290 Blanco, Ruthven, 19120, p. 231, F.M.N.H. ; San Francisco, ¥.M.N.HL.; Vera Cruz, Cope, 1887, p. 44, sae MENG pC 2AcS AGN Sakae Habitat and habits —V ery little is known of the habitat of this form and apparently nothing has been published concerning its habits. According to Gadow (1906, p. 277), “Guttatus does not leave the Tierra Galiente above the 3,000 feet level. Greater heights are an absolute barrier.” He (p. 325) considered the spotted phase as characteristic of the “open forests with dense undergrowth, or similar pitches of woodland, in the State of Vera Cruz and its confines with those of Oaxaca.” ‘That is, in the Atlantic Tierra Cahente, “ with its decidedly denser vegetation, with fewer decid- uous trees and much greater rainfall,” the “typical” spotted phase occurs. It was stated (p. 326) that the striped lizards prevail in the small open localities, and that “much tangled underbrush, broken terrain, well-wooded ravines, or meadows with tall grass and herbs, or rivers fringed with masses of shrubs,” were the features of the spots that yielded the most intermediate specimens. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS io Enemies —The Mexican whip snake, Masticophis mentovarius, was mentioned by Gadow (1908, p. 166) as preying upon these lizards. Affinities Because of its geographical and structural identity with deppii and its marked differences from gu/aris, its only other possible ancestor, guttatus is here considered as a direct derivative of deppii, or, perhaps more properly, of ancestral deppii-like stock. The occurrence of both guttatus and deppii in the same region is contrary to the usually applicable rule that directly related forms are not found in the same locality. This occurrence would seem, at first sight, to indicate speciation by mutation alone as suggested by Blanchard (1921, p. 253) for Lampropeltis californiae californiae which is found in southern California in coextensive distribution with Lampropeltis getulus boylii, from which it is apparently de- rived. For these Californian snakes, mutation alone, without isola- tion, seems to be the only possible means of speciation, but for the forms found in the region of Central America and southern Mexico, such as deppii and guttatus in this case, it is not necessary to resort eee eae Union Tsolation Union Migration and Intermingling Figure 17.—A THEORETICAL DIAGRAM OF THE MANNER IN WHICH TWO CLOSELY RE- LATED SPECIES, AT PRESENT IN CO-EXTENSIVE DISTRIBUTION, MAY HAVE EVOLVED FROM A COMMON STOCK THROUGH ISOLATION. D AND D’=DEPPIL; G=GUTTATUS wholly to this explanation. The theory of speciation by isolation (during which mutation has an oportunity to become of constant nature and hence of diagnostic value) may be used here. (See fig. 17.) It has generally been conceded by geologists (see Schuchert, 1929, p. 141) that the Tehuantepec region was lowered to below sea level in the post-Miocene or early Pliocene and that the resulting water barrier kept Central American forms isolated from those in southern Mexico for a significant length of time. For reasons to be presented in the general discussion at the end of this work it is presumed that the ancestral deppii stock became modified from the ancient /emniscatus stock during the late Miocene. Once differentiated, this generalized type spread northward and probably entered southern Mexico before the formation of the Tehuantepec portal. After the subsequent isolation and upon the re-formation of the Tehuantepec land bridge, later in the Pliocene, there is the possibility of a secondary union of the northern and southern faunal elements, 2307—31 6 74 BULLETIN 154, UNITED STATES NATIONAL MUSEUM Although guttatus is not found south of Mexico, deppii ranges south to Costa Rica, so during the submergence of the Tehuantepec region deppii is presumed to have remained unchanged or to have developed in Central America, whereas guttatus changed and evolved in southern Mexico. A secondary union of the land masses involved would have made it possible for the more plastic deppii to migrate northward into Mexico and not only to extend its range throughout the favorable area there occupied by guttatus but also to extend it northward along the Pacific slope, past the range of that form, into Colima and Nayarit as well. Such an explanation may also account for the great overlapping in the ranges of the closely related Ameiva undulata and Ameiva festiva which are also found in Central Amer- ica and Mexico. The foregoing diagram (fig. 17) is intended to illustrate the manner in which guttatus and deppii as closely related forms have assumed their present coextensive distribution. SUMMARY OF THE DEPPII GROUP The deppii group is confined to Mexico and Central America. Its representatives are alike in scutellation and resemble each other in color pattern, but the adult size is variable. There are three forms, deppti, cozumelus, and guttatus. In searching for the prototype among these one has to consider only deppii and guttatus since cozumelus is an insular form which has obviously been recently iso- lated and derived from deppi?. The remaining two forms are like- wise closely related, but deppii is thought to be the older unit because of its much closer geographical and colorational approach to lemni- seatus with which it agrees in size and from the stock of which the deppit group is obviously derived. Since a consideration of the genetic origin of the group as a whole, and hence of the prototypic depp, is given in the general discussion at the end of this work (pp. 251-260), the present consideration will be limited to the origin and relationships of the two derivatives, cozumelus and guttatus. Cozumelus is known only from the islands of Cozumel and Mujures off the coast of Quintana Roo in southern Mexico. These islands are very near to the mainland and the lizards are small like deppi, differing from it only in a slight but apparently constant modifica- tion of the color pattern. Because of this, and since guttatus and gularis, the other possible ancestors, differ greatly im size and scutel- lation, respectively, cozwmelus is thought to be a direct descendant from a common stock with deppi. The remaining form, guttatus, occurs only on the Mexican main- land. Because of its Salemne similarity to deppii and its obvious TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 75 differences from gulavis, it is considered to be a derivitive from the ancestral stock of the modern deppii. Its manner of origin is a matter of some doubt as indicated in the discussion of its affinities. Because of its coextensive distribution with the parent form, specia- tion by mutation, in place, is at first sight indicated. It is thought, however, that the natural geological changes that have occurred in \ \ Ne COZUMELUS--.-_---. @ 100 90 FIGURE 18.—MapP SHOWING DISTRIBUTION OF FORMS OF DEPPIL GROUP the region of Central America and Mexico, in this case at the isthmus of Tehuantepec, may have permitted speciation during a temporary period of isolation. Even this theory is thought to be in accordance with the principle of sveciation by mutation, which in itself repre- sents the sort of variation that is thought to give rise to new forms. For a discussion and diagram of the possibilities of this theory as apphed to deppit and guttatus see pages 73 and 74. 76 BULLETIN 154, UNITED STATES NATIONAL MUSEUM The apparent relationships within the deppii group are expressed by the following diagram (fig. 19). de ppi cozume lus Common stock of depp. and cozumelus guttatus Ancestral deppu stock FiGuRE 19.—DIAGRAM OF SUPPOSED RELATIONSHIPS WITHIN THE DEPPII GROUP THE SEXLINEATUS GROUP CNEMIDOPHORUS SEXLINEATUS SEXLINEATUS (Linnaeus) SIX-LINED RACE-RUNNER 1766. Lacerta sexlineata LinNarus, Syst. Nat., ed. 12, vol. 3, p. 364, (type locality, ‘‘ Carolina,” eastern United States).—GmeEtin, Syst. Nat., ed. 13, vol. 3, 1788, p. 105.—Daupin, Hist. Nat. Reptiles, vol. 3, 1802, p. 183.—SHaAw, Gen. Zool., Amph., vol. 3, 1802, p. 240.—Turton, English Ed. of Linneaus’ “Syst. Nat.,” vol. 1, 1806, p. 667.—FiTzincrer, Neue Classif. Reptilien, 1826, p. 51—HARLAN, Journ. Acad. Nat. Sci. Phila., vol. 6, 1827, pp. 18, 370. Scuinz, Naturg. u. Abbild. Reptilien, 1833, p. 102.—Hartan, Med. and Phys. Researches, 1835, pp. 144, 163.—SHERBoRN, Index Animalium (1758-1800), 1902, p. 897. 1774. Lacerta 6-lineata MUtirr, German trans. of Linneaus’ “Syst. Nat.,” ed. 12, vol. 3, p. 94, (all).—GarmaAn, Bull. Hssex Inst., vol. 16, 1884, p. 12 (part). 1789. Lacerta sex-lineata BONNATERRE, Erpetol., Tabl. Ency. Nature, p. 48.— SONNINI and LATREILLE, Hist. Nat. Reptiles, vol. 1, 1802, p. 245.—ScLATER, Reprint edition of Forrester’s ‘“‘Cat. of the Animals of N. Amer.,” 1882, p. 18. 1820. Teius lemniscatus MeRREM, Syst. Amph., p. 60, (part). 1820. Lacerta fallax Mrrrem, Syst. Amph., p. 63, (type locality, ‘ Carolina,” eastern United States). 1831. Ameiva sez-lineata Cuvier, Animal Kingdom, English transl. by H. M’Murtrie, Class III, Reptilia, vol. 2, p. 22—Horgprook, N. Amer. Herpetology, ed. 1, vol. 1, 1836, p. 63, pl. 6—BruHIN, Zool. Garten, vol. 15, 1874, p. 396.— NEHRLING, Zool. Garten, vol. 25, 1884, p. 259. 1831. Seps sealineata Cuvier, Animal Kingdom, English translation by H. M’Murtrie, Class III, Reptilia, vol. 2, p. 23. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 14: 1831. Ameiva sexrlineata Cuvier, Animal Kingdom, English translation by H. M’Murtrie, Class III, Reptilia, vol. 2, p. 28—Der Kay, Zool. of N. Y., vol. 1, pt. 3, p. 30.—Horeroox, N. Amer. Herpetology, ed. 2, vol. 1, 1842, p. 109, pl. 15; Reptiles, in White’s Statistics of Georgia, ‘‘ Flora and Fauna,” 1849, p. 13.—HALLOWELL, Reptiles, in Sitgreaves’ “ Rept. of an Expedition down the Zuni and Colo. Rivers,” 1853, p. 107. 1834. Lacertam sexlineatam WiEGMANN, Herpetologia Mexicana, p. 27 (foot- note). 1839. Cnemidophorus sexlineatus DumMERILT and Brsron, Erp. Gén., vol. 5, p. 1381.—F1TziNcrEr, Syst. Reptilium, 1843, p. 20—Gray, Cat. Liz. British Mus., 1845. p. 21—Bartrp and Gzirarp, Proc. Acad. Nat. Sci. Phila., 1852, p. 128.— HALLOWELL, Reptiles, in Sitgreaves’ “ Rept. of an Expedition down the Zuni and Colo. Rivers,” 1835, p. 145.—JANn, Indice, Sist. dei Rettili ed Anfibi Mus. Civ. di Milano, 1857, p. 40.—HALLOWELL, Reptiles, Pacific R. R. Sury. (Parke Route), vol. 10, pt. 6, sect. 2, no. 2, 1859, p. 28.—REINHARDT and LUr- KEN, Vidensk. Meddel. nat. Foren., 1861, p. 233; Vidensk. Meddel. nat. Foren., 1862, p. 168—GtnrHeEr, Zool. Rec. for 1865, vol. 2, p. 148; Zool. Rec. for 1866, vol. 3, 1867, p. 123.—PuHutipr1, Archiv Naturgesch., vol. 35, 1869, p. 42.—Perrers, Monatsber. Berlin Akad. Wiss., 1869, pp. 62-63.—CoUEs, Proce. Acad. Nat. Sci. Phila., 1871, p. 47.—Viranova Y Piers, La Creacion (Hist. Nat. ), vol. 5, 1874, p. 39.—W aRREN, Prelim. Rept. Expl. in Nebr. and Dakota in the years 1855-59, Reprint, Eng. Dept., U. S. Army, 1875.—JorpDan, Man. Vert. Northern U. S., ed. 1, 1876, p. 170.—Coprr, Proc. Amer. Philos. Soc. vol. 17, 1877, p. 65; Proc. Amer. Philos. Soc., vol. 17, 1877, p. 95—JorpAN, Man. Vert. Northern U. S., ed. 2, 1878, p. 170.—Corr, Bull. U. S. Nat. Mus., vol. 17, 1880, p. 18.—Cracin, Trans. Kan. Acad. Sci., vol. 7, 1881, p. 117.-— SmiTH, Rept. Geol. Sury. Ohio, vol. 4, 1882, p. 653.—Davis and Ricr, Bull. Chicago Acad. Sci., vol. 1, 1883, p. 31—GarMAN Mem. Mus. Comp. Zo6l., vol. 8, 1883, p. xiv—F1scHmr, Oster-Progr. Akad. Gymn. Hamburg, 1883, p. 2.— Hourrer, Cat. Reptiles and Batr. Coll. in Mo., 1883, p. 6 (printed privately) .— TrRuE, Vert. Animals of S. C., vol. 10, 18838, p. 237—Cracin, Bull. Washburn Lab. Nat. Hist., vol. 1, 1884, p. 8—Jorpan, Man. Vert. Northern U. S., ed. 4, 1884, p. 170.— BouLmnere, Cat. Liz. British Mus., vol. 2, 1885, p. 364.— HueHEsS, Bull. Brookville Soe. Nat. Hist., vol. 1, 1885, p. 41—Hay, Ann. Rept. Indiana St. Bd. Agri. for 1886, vol. 28, 1887, p. 214—BovtLencrEr, Zool. Rec. for 1886, vol. 28, ‘ Reptilia,’ 1887, p. 18; Cat. Liz. British Mus., vol. 3, 1887, p. 505; Ann. and Mag. Nat. Hist., ser. 6, vol. 1, 1888, p. 109.—KiIn«s- LEY, Riverside Nat. Hist., vol. 3, 1888, p. 482.—SrTesnecer, Proc. U. S. Nat. Mus., vol. 12, 1890, p. 644.—WerNER, Zool. Jalrb. (abt. Syst, Geog. u. Biol.), vol. 6, 1892, p. 200.—Corn, Trans. Amer. Philos. Soc., vol. 17, 1892, p. 42; Proe. Amer. Philos. Soc., vol. 30, 1892, p. 209.—GarMAN, Bull. Ill. St. Lab. Nat. Hist., vol. 3, 1892, p. 255.—Borrrerr, Kat. der Reptilien Samml. im Mus. Senckenb. Naturf. Ges. in Frankfurt, teil 1, 1893, p. 74.—WerRNER, Zool. Jahrb. (abt. Syst., Geog. u. Biol.), vol. 7, 1894, p. 383.—LornNBERG, Proc. U. S. Nat. Mus., vol. 17, 1894, p. 321—RuHoAps, Proc Acad. Nat. Sci. Phila., 1895, p. 386.—Corr, Amer. Nat., vol. 30, 1896, pp. 1008, 1011.—VAan DeEnBurRGH, Proc. Calif. Acad. Sci., ser. 2, vol. 6, 1896, p. 343; Proc. Amer. Philos. Soc., 1897, p. 4628.—Mocquarp, Nouv. Arch. Mus. Nat. Paris, ser. 4, vol. 1, 1899, p. 315.—JorpaNn, Man. Vert. Northern U. S., ed. 8, 1899, p. 201.— SMITH, Proc. Linn. Soc. New York, no. 11, p. 9—Bryer, Proc. La. Soe. Nat. Hist. for 1897-1899, 1900, p. 48——Gapow, Cambridge Nat. Hist., vol. 8, 1901, p. 549.—LAmpeE, Jahrb. Nassauisch. Ver. Naturkunde, vol. 54, 1901, p. 35.— STRECKER, Trans. Texas Acad. Sci., vol. 4, 1902, p. 97.—DLoENNBERG, Bihang 78 BULLETIN 154, UNITED STATES NATIONAL MUSEUM K. Svenska Vet.-Akad. Handl., vol. 28, 1902, p. 29.—Hay, Proc. Biol. Soc. Wash, vol. 15, 1902, p. 1834.—Gapow, Proe. Roy. Soe. London, vol. 72, 1903, pp. 112, 122—Coun, Zool. Anz., vol. 27, 1903, p. 187.—STonE and REHN, Proc. Acad. Nat. Sci. Phila., 1903, p. 33—Brimuiry, Amer. Nat., vol. 37, 1903, pp. 264— 265.—PRrRITcHETT, Biol. Bull., vol. 5, 1903, pp. 271, 284.—Morssr, Proc. Ohio St. Acad. Sci., vol. 4, 1904, p. 124.—HreNsHAw, Occas. Pap. Boston Soc. Nat. Hist., vol. 7, 1904, p. 5—Jorpan, Man. Vert. Animals Northern U. §., ed. 9, 1904, p. 201.—BouLeNncErR, Zool. Rec. for 1904, vol. 41, “ Reptilia,” 1905, p. 24— Bartey, N. Amer. Fauna, vol. 25, 1905, p. 44—HaArrMAN, Trans. Kan. Acad. Sci., vol. 20, 1906, p. 226.—Kinestry, Text Vert. Zool., 1906, p. 321.—FowtLrr, Proce. Acad. Nat. Sci. Phila., 1906, p. 111.—McAtrxr, Proc. Biol. Soe. Wash., vol. 20, 1907, p. 11—Drrmars, Reptile Book, 1907, p. 188.—BrimMirty, Journ. Elisha Mitchell Sci. Soce., vol. 23, 1907, p. 142—Brown, Proc. Acad. Nat. Sci. Phila., 1908, p. 118—Surracn, Zool. Bull. Div. Zool. Penn. Dept. Agri., vol. 5, 1908, p. 253.—Streckmr, Proc. Biol. Soc. Wash., vol. 21, 1908, p. 73.— HurrerR and Strecker, Trans. Acad. Sci. St. Louis, vol. 18, 1909, p. 23.— Hurrer, Trans. Acad. Sci. St. Louis, vol. 20, 1911, pp. 189-140.—WeERNER, Revision of “ Brehm’s Tierleben,” vol. 2, 1913, ». 148—BaArsourR, Mem. Mus. Comp. Zooél., vol. 44, 1914, p. 218—Barsour and Nose, Bull. Mus. Comp. Zool., vol. 59, 1915, p. 420—ANnon'ymous, Sci. Amer., vol. 112, 1915, p. 479.— STRECKER, Baylor Univ. Bull. vol. 18, 1915, p. 24—WkrienT and FUNK- HOUSER, Proc. Acad. Nat. Sci. Phila., 1915, p. 129——Brimuiny, Journ. Elisha Mitchell Sci. Soc., vol. 30, no. 4, p. 200.—DuNN, Copeia, no. 18, 1915, p. 6; Copeia, no. 53, 1918, p. 23.—DrEcKERT, Copeia, no. 54, 1918, p. 31—Hotr, Copeia, no. 72, 1919, p. 66.—DuNN Proe. Biol. Soc. Wash., vol. 33, 1920, p. 136.—Concrr, Key to Mich. Vert., Publ. by Mich. Agri. Coll., 1920, p. 41.— Bascock, Copeia, no. 85, 1920, p. 74.—Porrrer, Copeia, no. 86, 1920, p. 838.— BLANCHARD, Occas. Pap. Mus. Zool. Univ. Mich., no. 117, 1922, pp. 6—7.— Lopine, Pap. Alabama Mus. Nat. Hist., vol. 5, 1922, p. 24.—Srreckmr, Bull. Sci. Soe. San Antonio, vol. 4, 1922, p. 31—Camp, Bull. Amer. Mus. Nat. Hist., vol. 48, 1923, p. 410—SuHeERBoRN, Index Animalium (1801-1850), pt. 3, 1928, p. 506.—Over, Bull. 8. D. Geol. and Nat. Hist. Surv., vol. 12, 1928, pp 19-20. —HALLINAN, Copeia, no. 115, 1928, p. 19.—Vuiosca, Copeia, no. 115, 1923, p. 43.—WEED, Copeia, no. 116, 1923, p. 48.—Hott, Copeia, no. 186, 1924, p. 100.— StrecKER, Baylor Univ. Bull., no. 27, 1924, p. 37—FunNKHoOUSER, Wild Life in Kentucky, 1925, p. 77.—Forcn, Copeia, no. 141, 1925, p. 25.—Srrecker and Friprson, Contr. Baylor Univ. Mus., no. 5, 1926, p. 7—Srrecker, Contr. Bay- lor Univ. Mus., no. 3, 1926, p. 2; Contr. Baylor Univ. Mus., no. 6, 1926, p. 5; Idem, no. 7, 1926, p. 5—WRrRicHT, Hecology, vol. 7, 1926, pp. 883-84.—Viosca,. Heol- ogy, vol. 7, 1926, p. 310.—Brimiry, Journ. Elisha Mitchell Sci. Soc., vol. 42, 1926, p. 88.—WaArtTsoN, Naturalist’s Guide to the Americas, 1926, p. 484.-—HaAr- PER, JOHNSON and HOweE LL, Naturalist’s Guide to the Americas, 1926, p. 443.— Frison and Mirier, Naturalist’s Guide to the Americas, 1926, p. 471—Wotr- corr, Naturalist’s Guide to the Americas, 1926, p. 522.—Myerrs, Proc. Ind. Acad. Sci. for 1925, vol. 34, 1926, p. 286.—OrrenBURGHER, Proc. Okla. Acad. Sci., vol. 6, pt. 1, 1926, pp. 94-95; Copeia, no. 156, 1926, p. 145; Idem, no. 163, 1927, p. 47.—STREcCKER, Contr. Baylor Univ. Mus., no. 12, 1927, p. 14.—Cor- RINGTON, Copeia, no. 165, 1927, p. 100.—PiIckENS, Copeia, no. 165, 1927, p. 110.—Myers, Proc. Indiana Acad. Sci., vol. 36, 1927, p. 339.—Hast, Journ. Wash. Acad. Sci., vol. 17, 1927, p. 99—NosiE, Guide Leaflet Series, Amer. Mus. Nat. Hist., no. 69, 1927, p. 5.—Burr, Oceas. Pap. Mus. Zool. Univ. Mich., no. 189, 1927, p. 4; Journ. Kan. Ent. Soe., vol. 1, 1928, p. 58; Trans. Acad. Sci. St. Louis, vol. 26, 1928, p. 38; Key to Lizards of Kansas, printed TELMD LIZARDS OF THE GENUS CNEMIDOPHORUS 79 privately, 1928, p. 1—GtLoyp, Trans. Kan. Acad. Sci., vol. 31, 1928, p. 119.— Jones, Journ. Tenn. Acad. Sci., vol. 3, 1928, p. 12.—Strecker, Contr. Baylor Univ. Mus., no. 16, 1928, p. 12—OrTENBURGER, Mem. Uniy. Mich. Museums, vol. 1, 1928, p. 199.—Roppy, Reptiles of Penn., printed privately, 1928, p. 51.— SPRINGER, Copeia, no. 169, 1928, p. 103.—Forcr, Proc. Okla. Acad. Sci., vol. 8, 1928, p. 78.—PopE and Dickrnson, Bull. Public Mus. Milwaukee, no. 8, 1928, p. 43.—STRECKER and WILLIAMS, Contr. Baylor Univ. Mus., no. 17, 1928, p. 15.— ORTENBURGER, Copeia, no. 170, 1929, pp. 10, 27.—Jorpan, Man. Vert. Animals, ed. 13, 1929, p. 231.—Burr, Occas. Pap. Mus. Zool. Univ. Mich., no. 201, 1929, p. 8—StTREcKER, Cont. Baylor Univ. Mus., no. 19, 1929, pp. 5-6, 13.— CORRINGTON, Copeia, no. 172, 1929, p. 68—Nerrimne, Ann. Carnegie Mus., voi. 19, no. 3, 1930, p. 172.—Force, Copeia, no. 2, 1930, p. 28.—NosLe and TEALE, Copeia, no. 2, 1930, pp. 54-55.—OrTENBURGER, Copeia, no. 173, 1930, p. 94. 1851. Cnemidophorus serlineatus (part) DumMERILZ and DumeérIL., Cat. meth. Coll. Reptiles, Paris, p. 116—WestTepHAL-CASTELNAU, C. R. trav. Cong. sci. de France, tenu a Montpellier en dec. 1868, 1870, p. 19.—Bocourr, Miss. sci. Mex. et Amer. cent., vol. 3, 1874, p. 273, pl. 20c, fig. 11, 11a, pl. 20d, figs. 2, 2a—b.—Coprr, Bull. U. 8. Nat. Mus., no. 1, 1875, p. 45.— Yarrow, Bull. 24, U. S. Nat. Mus., 1882, pp. 8, 483.—GUnTuHER, Reptilia and Batr., in “ Biol. Centr.- Amer.,”” 1885, p. 25.—Patacky, Zool. Jahrb., abt. syst, geog, u. biol., vol. 12, 1899, pp. 253, 283.—Corr, Ann. Rept. U. S. Nat. Mus. for 1898, 1900, p. 593.— Gapbow, Proc. Zool. Soc. London, 1905, p. 216; Proce. Zool. Soc. London, 1906, p. 302.—ELLIs and HrenprERSoN, Univ. Colo. Studies, vol. 10, 1913, p. 78— STEINEGER and Barsour, Check List N. A. Amph. and Rept., 1917, p. 67; Check List N. A. Amph. and Rept., ed. 2, 1923, p. 73.—Pratr, Man. Vert. Animals U. S., 1928, p. 204.—CockERrELL, Zool. of Colo., Univ. Colo. Semicen- tennial Ser., vol. 3, 1927, p. 106. 1859. Cnemidophorus sex-lineatus Batrp, Reptiles, Pacific R. R. Sury. (Whipple and Ives Route), vol. 10, pt. 6, no. 4, p. 38——Hay, Ann. Rept. Indiana Dept. Geol. and Nat. Resources, vol. 17, 1898, p. 545-—-Hurrer, Trans. Acad. Sci. St. Louis, vol. 6, 1893, p. 259. 1874. Cnemidophorus sexrlineatus gularis (part) Bocourr, Miss. sci et Amer. cent., vol. 3, 1874, p. 278. 1884. Cnemidophorus tessellatus CRAGIN” Bull. Washburn Lab. Nat. Hist., vol. 1, 1885, p. 102. 1885. Cnemidophorus sexrlineatus HotpErR, Wood’s “ Animate Creation,’ revised and adapted to Amer. zool., New York, “ol. 3, p. 45. 1895. Cnemidophorus serlineilus Core, Proc. Acad. Nat. Sci. Phila., p. 387. 1899. Cnemidophorus sexlinsatus McLain, Notes on a coll. of reptiles made by C. J. Pierson at Fort Smith, Ark. (Contr. N. Amer. Herpetology), p. 2. 1900. Cnenidophorus sexlineatus serlineatus Corr, Ann. Rept. U. S. Nat. Mus. for 1898, 1900, pp. 598, 603-605 (part). 1902. Cnemydophorus serlineatus Toror, Nerthus, vol. 4, p. 201. 1903. Cnremidorphorus sexrlineatus serlineatus Brown, Proce. Acad. Nat. Sci. Phila., p. 546.—Burr and Burr, Amer. Mus. Novitates, no. 381, 1929, p. 9.— TAyYLor, Univ. Kansas Sci. Bull., vol. 19, 1929, p. 64. . Mexique General notes—During the early half of the past century, various workers, particularly those of Europe, have referred specimens from certain localities definitely outside of the range of sexlineatus to 2 This report is from McPherson County, Kans., and sezlineatus is the only form of Cnemidophorus inhabiting the region. In a recent letter Doctor Cragin writes that he feels that the identification is correct, but that the locality data may be in error. This seems very likely since the specimen was not personally collected by Doetor Cragin. SO BULLETIN 154, UNITED STATES NATIONAL MUSEUM this form. These errors have been amplified by certain later writers who have secured their information from the earlier publications. The most important point in the history of seawlineatus follow in chronological order. The 6-lined race runner was described by Linnaeus in 1766 as a species of Lacerta and between that date and 1839 it was referred to a number of genera. In 1836 Holbrook wrote of his “ Ameiva sex-lineata” as follows: “ This animal was certainly first described by Linnaeus * * * from a specimen sent him by Dr. Garden of Charleston, who furnished him with numerous rare specimens of plants and animals from Carolina. * * * It is impossible to understand what led him to consider this animal as the Lion Lizard of Catesby (vol. 2, pl. 68), with which it neither agrees in colour, habits, nor geographical distribution. * * * It is remarkable that most naturalists since Linnaeus have copied this error, and have given the same reference.” In 1839 Duméril and Bibron placed sealineatus in the genus Cnemidophorus and in 1848, Fitzinger (p. 20) designated this lizard as the type species of his genus Aspidoscelis, which however, has never been recognized. In their list of the specimens in the Paris Museum, Duméril and Duméril (1851, p. 116) recorded C. sexlineatus from the following localities: Savannah, Ga.; Vera Cruz, Mexico; Habana, Cuba; Charleston, S. C.; New Orleans, La.; and Martinique in the West Indies. The reports from the United States may have been correct, but the one from Vera Cruz may have been based on either gularis, deppii, or guttatus; the one from Habana, Cuba, was probably based on a specimen of Ameiva auberi; and, likewise, the one from “Martinique” must have been based on an Ameiva. Westphal- Castelnau (1870) apparently copied the errors of Duméril and Duméril, and in the same fashion Vilanova and Piera (1874) con- sidered this species as “inhabiting not only parts of the United States, but of Mexico and Martinique as well.” In 1878 Yarrow and Henshaw (p. 1640) reported sexlineatus from southern California on the basis of a series of specimens (U.S.N.M. No. 8630), and from Los Angeles, Calif., on the basis of one speci- men (U.S.N.M. No. 8631). Since both of these numbers were listed under tessellatus by Cope (1900), it is obvious that these reports were due to misidentifications. In 1866 Cope (p. 283) reported a series of specimens from near the city of Chihuahua, Mexico, with “six longitudinal stripes and unspotted interspaces” as C. sexlineatus sealineatus, but an examination shows that they were really inter- grades between gularis and perplevus, which are here recognized as subspecies of sevlineatus. Because of their close resemblance to each TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS Sl other at certain stages in the life history all three subspecies have been frequently confused as a glance at their respective tables of synonymy will show. Diagnosis —Structurally the 6-lined race runner is distinguished by its possession of four supraoculars, the absence of enlarged poly- gones or scutes on the postantebrachium, and its relatively small size. It retains a striped pattern throughout life and there are usually six, distinct, light lines on the back, although at times, especially in young examples, one or two, more or less faint and wavy, dorsal stripes may appear near the middorsal line. The third or upper lateral pair of stripes are placed closer to the stripes be- low them on each side than to each other and usually a more or less widened, yellowish, dull streak takes the place of a middorsal stripe. The ground color of the back varies, but it is usually brown, black- ish, or olivaceous; the upper sides are often decidedly lighter than the back in western specimens, in which case the two lateral fields are in sharp contrast on each side, the lower lateral remaining darker. The latter condition is not seen in gularis or perplexus. The under surfaces are usually white, although sometimes they stain to black- ish in preservatives, but they are never blue-black as in many speci- mens of gularis. Field spots are always absent in séalineatus. Description.—Snout blunt, rounded; nostril anterior to nasal su- ture; anterior nasal usually in contact with first upper labial, rarely with second; supraoculars normally 4; granules often not extend- ing past the posterior border of the third supraocular; frontopar- ietals usually 2; parietals normally 3; anterior gulars rather small, uniform, or with enlarged medio-lateral or median patches; posterior gulars smaller; mesoptychial scales moderate, largest cen- trally, larger rows 2-7; postmesoptychial granules fine, some us- ually on edge of posterior gular fold. Body moderately stout; ventral scutes arranged in 8 longitudinal and 28-38 transverse rows; dorsal granules moderately fine (east) to coarser (west); limbs well developed; brachials 4-8; antebra- chials 2-5; brachials and antebrachials usually continuous at a point of contact; postantebrachium with small or slightly enlarged gran- ules; femorals 4-8; tibials 2-4; femoral pores 12-30; tail elongate, tapering; caudal plates large, oblique, with moderate or weak longi- tudinal keels laterally. Coloration moderately distinctive, variable; ventral parts purplish, blue, greenish, yellowish, immaculate white or brownish; tail light below, and darker, usually olivaceous, above; dorsum lined at all ages; stripes normally 6, with the vestige of an additional pair usually showing anteriorly on each side in a line extending along the lower edge of the tympanum and ending abruptly behind it; middorsal 82 BULLETIN 154, UNITED STATES NATIONAL MUSEUM area usually with a more or less widened, dull, yellowish streak, but sometimes with an additional stripe or two; ground color of back variable, usually blackish, brown, gray or olivaceous; dorsal ground color sometimes in distinct contrast to lateral ground color; usually, however, area between dorsal stripes lighter on account of suffusion from the yollowish streak mentioned above; femora faintly reticu- lated with white or unicolor. A series of 1,522 specimens has been studied, but the data given below were taken from only 216 of them: Body, 27-82 mm.; tail, 45-164; total length, 72-246; width of head, 4.5-12.0; length of tail as percentage of total length, 55.5-72.1; width of head as percentage of body length, 11.1-17.7; supraoculars, 3 in 2 specimens, 4 in 205 specimens, and 5 in 9 specimens; supraocular granules extending forward to the middle of the fourth supraocular in 13 specimens, to the anterior border of the fourth supraocular in 165 specimens, and to the middle of the third supraocular in 388 specimens; fronto- parietals 2 in 195 specimens, 3 in 12 specimens, 4 in 7 specimens, and 5 in 2 specimens; preanals, 2 in 14 specimens, 3 in 102 specimens, 4 in 33 specimens, 5 in 3 specimens, 6 in 32 specimens, 7 in 6 specimens, 8 in 9 specimens, 9 in 8 specimens, 10 in 2 specimens, 11 in 2 speci- mens, 12 in 4 specimens, and 13 in one specimen. Cope (1900) mentioned that the three large anal plates are fused into one piece in two specimens from Florida (U.S.N.M. No. 15336), but that all other Florida specimens are “ normal.” Variation.—Perhaps the most significant variation found in sew- lineatus is geographical. Brown (1903, p. 546) recognized the close affinity with perplevus and wrote as follows: “ Two specimens from Pecos do not differ materially from eastern examples, except in hav- ing coarser dorsal scales, but four others, collected at Seymour, Tex., have similarly coarse scales. The pale dorsal area shows a tendency to contract and take on the form of a median stripe. All of the differences are in the direction of C. perplexus.” The dorsal scales are small and finely granular in examples from the Eastern United States, but they are usually enlarged in the area west of the Missis- sippi River and north of Texas. This coarse dorsal granulation in the west serves as an approach to perpleaus, as indicated by Brown. These subspecies are found to intergrade in the Panhandle district. Much individual variation exists in regard to these granules in all geographical areas and it seems that little more than an average difference exists between the eastern and western specimens in spite of appearances. The postantebrachium is usually covered by small granules, having characteristically but a small patch of shghtly enlarged ones. The patch may be so reduced as to become indistinguishable from the surrounding scales on the forearm or it may be so developed as to take on the character of small polygons. TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 83 At the point of intergradation with gularis there is a complete transition from the characteristic granules of sexlineatus to the iarger, better developed, polygons that are usually found in that form. The dorsal ground color is usually olivaceous, but in certain geographical areas, particularly the southern tip of Florida, the Tennessee region and Padre Island, off the east coast of Texas, the color becomes black and the rather characteristically widened, poorly defined, yellowish, middorsal streak becomes noticeably restricted and better defined so as to tend to produce the appearance of a moderately distinct middorsal stripe. Since this variation occurs in widely separated localities, it is not thought to have especial taxonomic significance. The variation in the position of the upper (third) laterals seems to have much more importance in this respect, since it is the approach of these two stripes toward the middorsal line that marks, in part, the transition to perplewus. This tendency and the fading and loss of the yellowish middorsal streak is clearly evident at the point of intergradation between the two. The ventral coloration of sexlineatus is like that of perplexus but differs from that of typical gularis in the absence of a deep bluish suffusion. Perfect transition from white to blue-black is seen at the point of intergradation between sex/ineatus and gularis in Texas and Oklahoma. Range.—This lizard is distributed over an unusually large area. It inhabits the lower levels in the Eastern United States from Mary- land west to the southern tip of Lake Michigan, northwest through southern Wisconsin to southwestern South Dakota, southwest to north central Colorado, south on the eastern side of the mountains to southern Colorado and the panhandles of Oklahoma and Texas, and southeast to the section of the Gulf coast lying just north of the mouth of the Rio Grande. A large number of records are available for sezlineatus, but there is a surprising lack of them from many areas. The reports will be presented below by States in an alphabetical series. ALABAMA.—Autauga County (Autaugavillie, U.S.N.M.). Bald- win County (General Report, Loding, 1922, p. 24). Calhoun County (Anniston, Dunn, 1920, p. 136). Cullman County (A? dell, U.S.N.M.). Greene County (Hutaw, Yarrow, 1882, p. 43, U.S.N. M.). Henry County (Abbevilie,U.S.N.M.). Jackson County (Sand Mountain, Holt, 1924, p. 100). Lee County (Auburn, U.S.N.M.). Lowndes County (Haynesville, U.S.N.M.). Mobile County (Mobile, Loding, 1922, p. 24, M.C.Z., A.N.S.P.). Montgomery County (Mont- gomery, Yarrow, 1882, p. 44, U.S.N.M.). Tuscaloosa County (Holz, Loding, 1922, p. 24, Mich.). 84 BULLETIN 154, UNITED STATES NATIONAL MUSEUM ARKANSAS.—Crawford County (Mulberry, K. U.). Garland County (Hot Springs, Strecker, 1924, p. 37, U.S.N.M., Baylor). Jefferson County (Pine Bluff, Hurter and Strecker, 1909, p. 23, U.S. N.M.). Lafayette County (Lewisville, K. U.). Lawrence County (Imboden, F.M.N.H.). Little River County (2 miles north of LAT ged LONGITUDE | RECORDS (COUNTY REPORTS) FOR CNEMIDOPHORUS SEXLINEATUS SEXLINEATUS LOCALITY THE SHOWING Map 9 FIGuRE Ashdown, Burt, A.M.N.H.). Pike County (3 miles south of Mena, Ortenburger, 1929, p. 10, Okla.). Polk County (3 miles south of Mena, Ortenburger, 1929, p. 10, Okla.; 4 méles west of Board Camp, Ortenburger, 1929, p. 10, Okla.). Prairie County (Devall Bluff, K. U.). Pulaski County (General Report, U.S.N.M.). Sebastian County (Fort Smith, Yarrow, 1882, p. 48, U.S.N.M., Stanford). TEIID LIZARDS OF THE GENUS CNEMIDOPHORUS 85 Washington County (/ayetteville, Hurter and Strecker, 1909, p. 23, .M.N.H.). COLORADO.—Adams County (Barr, Ellis and Henderson, 1915, p. 260). Denver County (Denver, Ellis and Henderson, 1913, p. 78). Jefferson County (Golden, Cary, 1911, p. 40). Larimer County (Arkins, U.S.N.M.). Las Animas County (Corrizo Creek, Ellis and Henderson, 1913, p. 78; Ponia, near Trinidad, Ellis and Hen- derson, 1913, p. 78). Prowers County (Zamar, Mich.). Washington County (Akron, Mich.). Weld County (@reeley, Ellis and Hender- son, 1913, p. 78.).. Yuma County (Wray, Ellis and Henderson, 1913, p:. (8, A.M.N.H.). CONNECTICUT.—Yarrow (1882, p. 44) reported a specimen from Middletown, Middlesex County, but this record does not seem to be backed by a specimen in the United States National Museum at the present time. Babcock (1920, p. 74) has written as follows: “ The striped lizard reported by E. Smith (1899) as a ‘hearsay’ record from Connecticut occurs normally from Maryland to Florida.” DISTRICT OF COLUMBIA.— (Terra Cotta, East, 1927, p. 399, U.S.N.M.). FLORIDA.—Alachua County (Gainesville, A.M.N.H., Maich.; Micanopy, Yarrow, 1882, p. 43, U.S.N.M.). Brevard County (Can- averal, A.M.N.H.; Faw Gallte, A.M.N.H., M.C.Z.; Fort Lauderdale, U.S.N.M.; Georgiana, Yarrow, 1882, p. 44, U.S.N.M.; Indian River City, U.S.N.M., Mich.). Broward County (Hallandale, M.C.Z.; Pompano, M.C.Z.). Dade County (Miami, A.N.S.P.). Duval County (Arlington, Yarrow, 1882, p. 44, U.S.N.M., M.C.Z.; Cedar Creck, Hallinan, 1923, p. 19; Hastport, A.M.N.H.; Jacksonville, Deckert, 1918, p. 31, A.M.N.H.). Escambia County (Pensacola, Yar- row, 1882, p. 48, U.S.N.M., M.C.Z., A.N.S.P.). Franklin County (St. Vineent’s Sound, A.M.N.H.). Jefferson County (Lake Miccosu- lee, F.M.N.H.). Lake County (Lake Harris, Cope, 1900, p. 597, U.S.N.M.). Lee County (Fort Myers, A.M.N.H.; Labelle, Mich.). Manatee County (Hgmont Key, F.M.N.H.; Lemon City, U.S.N.M.; Little Sarasota Bay, Yarrow, 1882, p. 44; Miakka, U.S.N.M.). Marion County (General Report, F.M.N.H., Carnegie; Hureka, A.M.N.H., Mich.). Monroe County (Big Pine Key, Fowler, 1906, p. 111; Boca Chica Key, Fowler, 1906, p. 111; Grassy Key, Fowler, 1906, p. 111; Indian Key, Yarrow, 1882, p. 43; Key Vacas, Fowler, L0G ep. Ail: Key West, Cope, 1900, p2e597, . W:S.N.M.,. M.C:Z,, F.M.N.H., C.A.8., A.N.S.P.; Anights Key, Fowler, 1906, p. 111; New Found Harbor, second key southwest of Big Pine Key, U.S.N.M.). Orange County (Oakland, Loennberg, 1894, p. 321; Or- lando, Loennberg, 1894, p. 321). Palm Beach County (Lake Worth. U.S.N.M.; West Palm Beach, M.C.Z.). Pasco County (Argo, A.N.S.P.). Pinellas County (Clearwater, Yarrow, 1882, p. 44. S6 BULLETIN 154, UNITED STATES NATIONAL MUSEUM U.S.N.M.; Point Pinellas, U.S.N.M.; Seminole, Mich.; Seven Oaks, A.M.N.H.). Polk County (Auburndale, U.S.N.M.; Eustis Lake, U.S.N.M.). Santa Rosa County (Santa Rosa Island, U.S.N.M.). St. Lucie County (Sebastian, M.C.Z.). Volusia County (Hnterprise, Mich.; Volusia, Cope, 1887, p. 65, A.N.S.P.). Wakulla County (Gulf Coast south of Tallahassee, ¥.M.N.H.). GEORGIA.—Baldwin County (Milledgeville, Yarrow, 1882, p. 48, U.S.N.M., M.C.Z.). Berrien County (Nashwille, Yarrow, 1882, p. 44, U.S.N.M.). Camden County (St. Afarys, Cope, 1900, p. 597, U.S.N.M.). Charlton County (Billy’s Island, Honey Island, and Minne Lake Island, Okefinokee Swamp, Wright and Funkhouser, 1915, p. 129). Chatham County (Savannah, Cope, 1900, p. 597, U.S.N.M., M.C.Z.). Grady County (Beachton, F.M.N.H.). Turner County (Ashburn, A.M.N.H.). ILLINOIS.—Hardin County (Garman, 1892, p. 256. listed an ancertain record from “ Cave in Rock,” which is not to be accepted until verified). Henderson County (General Report, U.S.N.M.). Kankakee County (Z//inoi, F.M.N.H.). La Salle County (Ottawa, Garman, 1892, p. 256). Marshall County (Henry, Garman, 1892, p. 256). Mason County (Topeka, Iowa State). Morgan County (Meredosia, Weed, 1923, p. 48). St. Clair County (Bluffs near Bluff Lake, Baie 1893, p. 259). INDIANA.—Frankln County (General Report, Hughes, 1885, p. 41, “ I have seen only one of these specimens from the county ”). Knox County (Hay, 1893, p. 545, “ This species is included in the fauna of Indiana on the testimony of Mr. Robert Ridgway of the National Museum. He writes that while he was collecting snakes at Monteur’s Pond, near Wheatland, he climbed a buttonbush to snare a big water snake. While thus engaged he saw a specimen of what he is positive is this species, with the appearance of which he was previously very familiar.” This record served as the basis of Myers’s, 1926, p. 286, report of sealineatus from Indiana. =a eee Bi 42, 46, 53, 60, 65, 72, 83, 113, 133, 142, 177, 202, 204, 207, 210, 212, 215, 218, 220, 236, 242, 243, 246. DistrictotiC oluMibla sae eee 85 ID OGOENG > ees ok ee ee OD LOD ir owming 25 = 2.2 Se eee 194 DiutehwGuian as ee eee 39 Duteh eeward Tslands-—-=-=_-222--=-==2 42, 50, 53 CVINOS MUNG a= oe ne eee Ona ZOD Wer-lavimcen shee eee ors 95, 119, 139, 196, 197 Has) ae... 3 See a ee 94, 95, 119, 196 TET TiO ee ee ee 94, 196 Enemiess= =. -- aoe 62, 73, 95, 120, 139, 197, 221, 240 Environment, influence on coloration-_------ Lis 69, 72, 153, 154, 155, 156, 157, 166, 169, 170, 171, 190, 218. influence on speciation. _——--- 5 HERES DONO ULE a ee 251 Meeding activa ty. a2 = ee 92, 93, 94, 119, 139, 192, 194, 195, 212 Bloridae 2-422 =e ee ee 85 HOO =: ee 93, 94, 119, 139, 194, 195, 212 OSSIIStermentLONM Ofna a eee 252 French Guiana a2. 2.8. Se Se ee ee 39 General discussion of relationships----------- 251 INDEX 285 Page Page Generic integradation__________- 12S pol Poe) eb4 «|, Totelapaces ===. = 4 a 5, 69, 102 ReeO Praia TAChS os ae ee ae ne 3,4,5 | Locality records, methods of presenting_____- 6 ‘Geological changes, influence of_____----_-__- 4,5 bablesdh: = 2e5 ate Ae ee) side REST) NaI eee ee yh Pee ee ee Ee ee 86 42, 46, 49, 53, 60, 65, 72, 83, 113, MMmang win when MZard 2-2 Sos ee 218 133, 142, 177, 202, 204, 207, 210, OG (ae De eS SS ee ee 39, 61, 117 212, 215, 218, 220, 236, 242, 243, Grular region, figure of_....-...-.......-..-=- 24 236. Strom @ahifornia, Tslands. 3 2- -. 5. 8... PSRE 1 CUVOUISTAIVAS 2 = aoa OE SE ee Se eee 88 204, 205, 207, 210, 212, 215, 218, 242, 243, 246 bower-Cahifomiiia _. = -=2-52-5- 5222 187, 202, 220, 238 Ma EILU TIC US ee oe eee 251 | Many lined race-runner. -...-.._.-.-=.--L-=. 56 Aue en ee Se eee eee 19; .'|) IMlaps,-growps_..----22-...- 54, 75, 145, 222, 248, 259 39, 62, 72, 91, 118, 1387, 188, 205, 207, 210, 212, species____-_ 38, 41, 61, 71, 84, 113, 133, 178, 237 218, 220, 239 Slatin amen a ee = he ee ee Pee 39 Wabitss—.-- 19) '99°5062,'92: 118; Tas: 101, 2197290939 | Marland. ....-.2.---- 22-22. eee ee 88 Habits, ontogenetic changes in__--__-.______- 118 | Masticophis flagellum flavigularis____________- 126 mieadapieaices: terres Of: 92 = as 22, 23 A MOMURLILS to ee ae 197, 221 LE LALECU UES ae ene a a ee emo 251 ICES Ga tac eee seers 240 SS ELH ENT ATCO 1 Ue hs Tee 92 MLCNEORAMIAGG oF oe oe ee 62, 73 BABSLOLICH CO WIGW a2 ose ee ee ee oe 8 taeniatus taeniatus_——— —- 197 Holbrookia maculata approximans_-_-----_--- W338), | Miatingudaves: 2 ==- 3) e230 ee 196, 239 LOTR eed eek ne Se URS FUL) eave AtrSt tne ee ee on eae 3, 154, 155, 156, 157 ners ae ee ee ee ee ee OU GI! |) Wlexican Tacenumners=--= 22> -. 2 ee 66 inner yunnis Group: ===. === 22-522 se Ss 16, | Mexico____- 19,61, 65, 72, 115, 1386, 187, 249, 250, 251 20, 140,141, 143, 226, 246, 253, 254, 257, 258, 259, 260 VICT ORION ANS tate = eee a ee ane ee 252 pt ren tI mene epee te ae oe ae ees bee ee SS SS 184s RAVI CeTIOl sek see ea See ee eee 73, 225, 248,259 MA ACen ere eee IS SEE ee ee 252k PeMLISSISSLD Des == 2-2 eben. so a ee eee penne 88 JANE GES Sat ee a le ee ee 86 in @AVINSSOMTIe Se sea = ee ee se nee ae 88 Meri uen eTIOd = = = = ae 95 19697. ||) tonttoniWalenstnigusnea = ea es ee 11 rans oe ee ee 86 | Monserrate Island race-runner____--_____-_- 242 Intergradation, evidences of.________-------- 12; | Museums; abbreviations for__--=_---_______- 8 13,/42) 53, 65,82, ‘83, 96, 120, 121, | Mutations, importance of__.__-__---..-____- Baie 139, 199, 200, 204, 205, 208, 253, Nebraskans Sts. Seo She 2 cee ee a 88 254. INEUSTICUTIS Epa ee ae ee Ee 252, 253 method ofdetermimnation=—-= 3,455: | Nevada. = --=2..- =. =.=. 2 222 2 185 SEEN LAG G TOT 8 pe) ee re es See oe, De ee we IMIGK CG Se a5 a cee toe a ee EF 135, 185 ah een Se eee a Oe ron SGle | eee ara Uae ee Se et Eo as Oe ee 39, 62 LLB VTS EE A ST oe Se 252)9| OTE ne GOaroninea-== == =. ee oe ee ee 88 MS PUTTS ACE LEY lane ee ee ee SONG5F | MOkiahe metas ane See ee 89, 114 GuiltoiCaliformia = =-2=— == 202, 220, 238, 239 | Old Providence Island. -_-.-----.-__-___..- 39 DRSOT Te ess sea eee ee TSR 2022381239) || sOnemidophonus.--- = ee! Se ee 11 Tsplanion, imntuence or. =-=--=.2=-_----—=- 4, 5, 73, 110 EDL MENINGES AE 7 preservation of primitive types by - 56 SCINEGON ae = ee 151 ESRI See ee eo ma Se rie ten ee 2 De Si, || wOnhionomon =: s2- 2e = es ee ee ee ee 251 LEGS UTE SS SS OO i Be 2D 2 PONCE OM Sate cee ee Syne ee Bee 186 PERL TUU (ALTE een a ee 21 NEF ET NY ee gel oe st le ae ee en eee 39 RTT aViee eee teas se ee ee ee RSt || Wasa Peano oe Noe a ee ere 46 Pe veLO aleO PeNetde = ae n= ho 8 soe Soe ae 14. | Penial characters. =< --- 22 <-.2 sense ae ea 20, | Relationships, diagrams of__-.-.-.----------- 56, 30, 40, 53, 54, 253, 254, 255, 257, 259, 260 76, 140, 146, 199, 225, 249, 253, 254 TODOS OTIS oe a ee eee ee 252 | Richardson’s law, remark upon-.------------ 175 GOALS Olea ee = 2 oe te eee 94-139 3196's DOR ORG -MUNTOR 1222 e eee eae eee 197 286 INDEX Page Page Running movement__-_-_ 39, 93, 119, 192, 193, 221, 240 PONS ps Anes oak Sasa Pe ee ee 10 Salvad ors a= = ieee See ee ee eee 62 OMe: = 5. eee ee eee 11 Salvadora grahamiae heralepis________________ 139 CYONCUS— sass re = oe ne eee 11 San Jose Island race-runner____--_-------___- 244 ocellijers. i one eee 9, 11, 43: San Jose Island whiptail-_.-_.. _.-_--._-.-. 202 Ent ENR CUS a en ee 11 San Pedro Martir Island whiptail____._____- 20 Sila enn CSSeC eas eee Sie eee ae ee 90 San Pedro Nolasco Island spotted whiptail_. 211 | Tessellated lizard__........_..._...-_________- 146 Santa Catalina Island spotted whiptail______ 213i eeessellavusnCrolp sae = ee ee 14, 20, Sceloporus undulatus undulatus. _-_-_-_-__-_- 94 120, 143, 146, 250, 253, 254, 255, 257, 258, 259, 260 WOOLECOSCUT US ae oe ee ee eee Spill hexase se ie oan ene ee 90, 114, 136, 186 SOD Soe Se Re Bee ee ae eae 8, 10 EAUSYCUCILC IS eae = ee eee oot ae 30 COENEN GS cobain ot as Soe ee 30, 32 Ocelltjen jae oe eee 43 lemmiscnive: == eee ore 305 Rigerligardt-= <2 a2 oie aes ee oe Ue 146 TULL UTS pre Ne ee oe See ees Se O46 sae | Oba SOs aes ae eee oe eee eee Cees 39 ET LENCO LORE es eee oie ae ee RO tee OT CG Sen aes oe ee es ae 11, 12, 13, 14, 251, 254 sexlineatus eroup=-_- >see a 2ORAG 120) 3: | WD AGRGoe = mee = See eo ey eee ee 191 TAO 43 VAST 250 Ae da ros. OO eo te LOO e200. Oem ee NEL CLA Cle = Sek nesters eee eee aa 39 Sexual dimorphism_-__---_-____- ne ae 37, 49 AUDINOMOIS: ee ae ee ae 252, 253 Six-lined nace-rinner ss ee f6) Urinary, bladder, absenceiofe=-o- =e seen 18 ey Ren ha fae ee in ee re ee ee 18) ah aUba ey keee oe ee el oa eee ea el 136, 186 SMa Ss suSedias food sas ae ee O4F AVAL Ati OT COLOLA D1 Om a] a= ere eae 3, 4,9, SONOLAMTACe-nlInne na ee eee 122 16, 17, 18, 21, 42, 60, 64, 68, 69, 132, 153, Sonphe@ anol s eat eee eee eee ae 89 154, 155, 156, 157, 174, 210, 217, 235, 236. SOUbhDakotassat ose eee ae eee ee 90 COLT DCH) eae eee are ee 3, 4, 9, 21, SDeOCiaO Or ee a ee, Gt a eS Aa 42, 83, 101, 102, 112, 132, 174, 235, 236 Species concept Olas s > ha ee a ee 3 jolanrccy(oy Koysavepil os ss ee 5 DSU] Ais Sac er Renee ee gee, eee ee 3, 4,5 StRUCULITal seas eee Sy le nelisoe of uncertain taxonomic position —____ 249 49, 60, 65, 71, 112, 175, 176, 233, 234, 235, 236 remarks on lumping of_________-____- 3.) MENCZUClAs. -- =n ea NE ae ae 39, 50 remarks on retention of indefinable - BP |) e MeneZ el arielS len Seen ee ee 39, 42 Speedics 22 2h- 2 «28 a ee 119, 138, 192, 193, 220, 239 Wenicanian.. 2 a ee re 11, 14 Spottedmrace-runnere se ee ee 97 beldingi_____- 141, 226, 227, 228, 229, 230, 232 Stomachicontentsas.= sae 94, 139, 195, 196 caenuleds-~2 <2 ee eee 232, 240 SiDSPeClesCONGeDt Ola ee 3, 158 cenalbensis== =" 2s =e ee ee 14 DUMIMaArye Geppil erOU pees. ee 74 espiritensis._.___ 228, 230, 231, 232; 233, 239 hyperythrusieroup=—— 246 TROMCISCENSS ee 228, 230, 232, 233, 234 leminiscatusierowpes-= 2-2) eee 53 hyperythra____ 14, 226, 228, 229, 230, 232, 235 Sexlineatus group =-- 22 = - 144 beldingi___ 227, 231, 235, 236, 239 tessellatus croup ease == ee 221 DELO 227 Sunning and need of warmth-_-_-__-_---_____- 192 hyperythra_ 226, 231, 232, 236,239 Sate S anid sae ee Pas SER tae ae tego el 39 Schinidiiee= ===) eee 228, SVSPCMMIALIC) NOLESs= se ee ee OD ASAT DE Ols 230, 231, 233, 234, 236 79, 101, 125, 152, 199, 205, 208, 211, 213, 228, 249 Dicie=—— =e ee ee 242 Tantilla nelsoni____________ pv ote eae Re 106 SeTICEUS 2 teen ee ee 231, 232, 244 Mara Cina ae eee eee lee ee oe a ee ee AGis || VArginiges coe se oo eee 91 Taxonomic refinements, reference to_________ 10 | Water, concealment in___..._...------- 119, 138, 194 AGYe\f) ON ae ee ie i See a ads he te ee 18 | Western orange-throated race-runner--—-_----- 226 Teiidae, discussion of phylogeny of___-_ 251, 252,253 | Western reticulated whiptail_-__.--__-----_-- 208 REVUS eee eee ee eee. eee ee Qe ONAS 25s WHASCONSINe nee ose ee ee er 91 COLETLLLCOCE DIL Ce na eee 31, 32 CUCTICUS Rose nee ee er aD 30, 32, 46 LOUIS COM ULS tree cert ta eee ee aes 30, 76 C) SMITHSONIAN INSTITUTION L “MII IBRARIES Ui] 3 9088 01421 2354 | MT