Bulletin of the Chicago Academy of Sciences, Vol. 10, No. 11
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Vol. 10 No. 11 Bulletin of the Chicago Academy of Sciences A Solution to the Long Neglected Holbrookia lacerata Problem, and the Description of Two New Subspecies of Holbrookia Ralph W. Axtell University of Texas, Austin Chicago Published by the Academy 1956 The Bulletin of the Chicago Academy of Sciences was initiated in 1883 and volumes 1 to 4 were published prior to June, 1913. During the following twenty-year period it was not issued. Volumes 1, 2, and 4 contain technical or semi- technical papers on various subjects in the natural sciences. Volume 3 contains museum reports, descriptions of museum exhibits, and announcements. Publication of the Bulletin was resumed in 1934 with Volume 5. This series is now regarded as an outlet for short to moderate-sized original papers on natural history, in its broad sense, by members of the museum staff, members of the Academy, and for papers by other authors which are based in considerable part upon the collections of the Academy. It is edited by the Director of the Academy with the assistance of a committee from the Board of Scientific Governors. The separate numbers, issued at irregular intervals, are distributed to libraries and scientific organizations and to specialists with whom the Academy maintains exchanges. A reserve is set aside for future need as exchanges and the remainder of the edition offered for sale at a nominal price. When a sufficient number of pages have been printed to form a volume of convenient size, a title page, table of contents, and index are supplied to libraries and institutions which receive the entire series. Howard K. Gloyd, Director. 163 3 Bulletin of The Chicago Academy of Sciences Vol. 10 Bulletin of the Chicago Academy of Sciences A Solution to the Long Neglected Holbrookia lacerata Problem, and the Description of Two New Subspecies of Holbrookia Ralph W. Axtell University of Texas, Austin An interest in the lizard genus Holbrookia that dates back to my high school days in southern Texas has burgeoned to the point where a monographic treatment of the group is now in preparation. During the earlier phases of research with the Texas members of the genus, a taxonomic problem came to light which I feel should be published preliminary to the full report. This problem, and the concomitant description of two new subspecies which are a result of the study, is to be presented here. Without the cooperation and assistance of many of my friends and associates an undertaking such as this could never have been realized. I am wholeheartedly grateful to the following people and institutions. For the many courtesies afforded me by the persons in charge of the collections at various institutions, and for loan of specimens, I would like to thank Dr. Doris M. Cochran of the U. S. National Museum ( USNM), Dr. Charles M. Bogert and Bessie M. Hecht of the American Museum of Natural History (AMNH ), Dr. Bryce C. Brown of the Baylor University Museum (BUM) and for the loan of his private collection ( BCB), Dr. W. B. Davis of Texas A. and M. College (Texas Cooperative Wildlife Collection—TCWC), Dr. Carl D. Riggs and Dr. Charles C. Carpenter of the University of Oklahoma Museum of Zoology ( UOMZ), Dr. Edward H. Taylor and John M. Legler of the University of Kansas Museum of Natural History (UKMNH), and Dr. Norman Hartweg, Dr. William E. Deullman and Jack Damon of the University of Michigan Museum of Zoology (UMMZ). For gifts or loans of specimens from individuals, I would like to thank M. J. 4 Bulletin of The Chicago Academy of Sciences Vol. 10 Fouquette, Dr. J. S. Mecham, Dorothy Trevino Robinson, George G. Henderson, John E. Werler, Michael Robinson, and Dr. Pauline James. Specimens in my personal collection are marked RWA. I wish especially to thank Dr. Marshall C. Johnston, James R. Tamsitt, and Dr. Henry H. Hildebrand for their companionship and assistance in the field. Mr. Tamsitt deserves special mention for securing and observing both Holbrookia lacerata and Holbrookia maculata in the zone of sympatric overlap near Big Spring, Texas. Credit for the photographs should go to Isabelle and Roger Conant, who graciously lent their time and ability while snowed under with work on the forthcoming Field Guide. Mr. Conant has carefully read the manuscript and offered many helpful suggestions. Finally, I thank Dr. W. F. Blair and the Texas Natural History Collection (TNHC) of the University of Texas for making available specimens, space and material for carrying on my research. Dr. Blair has read the paper and offered valuable suggestions and criticisms. HISTORY OF THE PROBLEM The problem is an old one, dating back to the description of Hol- brookia lacerata by Cope (1880). At that time, Cope supplied an excellent description of the material he had at hand, but included along with his definition, locality "records" from two rather widely separated areas ( about 120 airline miles) in Texas. Quoting from Cope : "The most northerly locality for Holbrookia lacera [lacerata] with which I am acquainted is in Erath County, west of the upper Brazos. Mr. Boll found it rather abundantly there and in Comanche County. Southward it has been found by Mr. Marnock on the Guadalupe River in Kendall or Comal County." It should be pointed out that Cope cited no definite locality records, nor did he indicate whether he possessed specimens from either of the two regions mentioned in the quotation above. I have been unable to find specimens antedating Copes work from either of these areas. I did find two specimens in the U. S. National Museum, however, which are marked as types. One of these (USNM 10160A) a male, fits Copes description almost perfectly. These two cotypes were collected by G. W. Marnock, but were reported from Helotes, Texas, which is not on the Guadalupe River. It is possible that Marnock might have collected the cotypes on the Guadalupe River, and shipped the specimens from Helotes without locality data. Shipping points rather than actual locality records have the notorious habit of slipping into museum catalogs. In this case little harm is done because Helotes happens to be in, or at least very near, the known range of the lizard described. 163 1956 Axtell: New Subspecies of Holbrookia 165 Copes use of plural locality data served as the initial point of confusion regarding the interspecific relationships of the lizard he was describing. By citing information for two well-separated localities, Cope, as will be shown presently, innocently included two distinct species (H. lacerata and H. maculata) under the single specific name Holbrookia lacerata. Several papers subsequently were published which tended to perpetuate the initial confusion instead of alleviating it. Stejneger (1890) reduced the newly described H. lacerata to a subspecies of Holbrookia maculata, and was the first and only worker to point out the identity of the types as associated with the Helotes type locality. Although Stejneger recognized only one type locality, he continued to lump the type specimens from Helotes with the specimens from north central Texas, just as Cope had originally done. Cope ( 1900), himself adopted Stejnegers views and treated lacerata as a subspecies of maculata. Schmidt, who published the first comprehensive review of the genus Holbrookia (1922), enumerated the difficulties encountered in distinguishing certain H. lacerata from H. maculata. He recognized several of the paramount characteristics of H. lacerata, such as the distinctive dorsal coloration and markings, the subcaudal black spots, and the variable lateroventral black spots, but he regarded these characters as unstable since some of the material he examined did not exhibit them. Although Schmidt listed "authentic" localities1 in seven Texas counties ( Crockett, Kerr, Kendall, Bexar, McLennan, Erath and Comanche), he failed to notice any geographic discontinuity in the incidence of subcaudal spotting. I will show that such discontinuity does exist. Schmidt apparently was not swayed by the changing views of his predecessors with regard to the subspecific status of H. lacerata, because he continued to recognize it as a full species. In the latest checklist (1953), this point of view is retained, but the original double type locality is also retained. General works, mostly checklists, regional lists, and handbooks which have appeared since 1922, have tended to follow the views of Stejneger (1890) by considering lacerata a subspecies of H. maculata. Smith (1946) included a section on Holbrookia which is perhaps the most detailed treatment of the genus thus far. Smith employed the original 1 Schmidts use of the term "authentic" is ambiguous, as he was able to examine only six specimens. These six specimens could have hardly come from seven different counties. The word authentic implies first hand identification by the investigator, not the allocation of literature records. 6 Bulletin of The Chicago Academy of Sciences Vol. 10 1956 Axtell: New Subspecies of Holbrookia 167 dual type localities for lacerata and recognized it as a subspecies of Holbrookia maculata. DISCUSSION AND DESCRIPTIONS Now that a considerable quantity of material is available from a large number of localities, it can be shown that the Holbrookia lacerata of earlier workers consists of two distinct species. These are : Holbrookia lacerata, which has very distinctive subcaudal spots, a vivid dorsal pattern of light-margined dark blotches arranged on a grayish-brown ground color, and inhabits the Edwards Plateau of Texas and regions to the southward. Holbrookia maculata, which has no trace of subcaudal spotting (H. m. dickersonae sometimes has dark pigmented bands on the subcaudal surface), a vivid or obliterated dorsal pattern of dark blotches with light posterior margins on a tan, brown or gray ground color, and is found north and west of the Edwards Plateau, and on the plains of the Mexican tableland.