?>'ajPr-(\)Cx^ Or\?D^S3 /iPR 80 1971 -tA^U V VOLUME 16 1969-1970 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in chapters I and II (only) of the Style Manual for Biological Journals, 2nd ed., published in 1964 by the American Institute of Biological Sciences, Washington, D. C. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. Illustrations should be proportioned for one or two column width reproductions and should allow for insertion of legend if occupying a whole page. An abstract not exceeding three percent of the length of the original article must ac- company each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. The editors also recognize the policy adopted by the Federal Council for Science and Technology, and endorsed by the Conference of Biological Editors, that page charges for publication of scientific research results in scientific journals will be budgeted for and paid as a necessary part of research costs under Federal grants and contracts. Accord- ingly, writers crediting research grant support in their contributions will be requested to defray publication costs if allowable under the terms of their specific awards. Illustrations and tabular matter in excess of 20 percent of the total number of pages may be charged to the author; this charge is subject to negotiation. Exchanges are invited from institutions publishing comparable series but subscriptions are available if no exchange agreement can be effected. Separate numbers or volumes can be purchased by individuals and subscriptions are accepted. Remittance should accompany orders from individuals. Authors may obtain separates of their articles at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to Dr. Royal D. Suttkus, Department of Biology, Tulane University, New Orleans, Louisiana 70118. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. and Bat. Price per number (flat rate) : $1.50 Price per volume (flat rate) : $5.00 Meade Natural History Library Tulane University Riverside Research Laboratories Route l,Box46-B Belle Chasse, Louisiana 70037 Gerald E. Gunning, Editor Alfred E. Smalley, Associate Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Vickie L. Haddenhorst Assistant to the Editors CONTENTS OF VOLUME 16 Number P^ge 1. VARIATION AND RELATIONSHIP OF THE STUDFISHES FUNDULUS CATENATUS AND FUNDULUS STELLIFER (CYPRINODONTIDAE PIS- CES) ; Jamie E. Thomerson 1 SWINGLEUS POLYCLITHOROIDES GEN. ET SP. N. (MONOGENEA: GYRODACTYLIDAE ) FROM FUNDULUS GRANDIS BAIRD AND GIR- ARD Wilmer A. Rogers 22 A NEW TREMATODE, LEPIDODIDYMOCYSTIS IRWINI N. G, N. SP. (DIDYMOZOIDAE) FROM A MARINE FISH, MENTICIRRHUS NASUS Satyu Yamaguti and Shunya Kamegai 26 2. THE COACHWHIP SNAKE, MASTICOPHIS FLAGELLUM (SHAW): TAXONOMY AND DISTRIBUTION Larry David Wilson 31 3. INTRODUCED GULF COAST MOLLUSCS Dee Saunders Dundee 101 TYPE SPECIMENS OF FISHES IN THE TULANE UNIVERSITY COLLEC- TION WITH A BRIEF HISTORY OF THE COLLECTION Royal D. Suttkus ll6 4. A SYSTEMATIC REVIEW OF U ROM ACER CATESBYl SCHLEGEL (SER- PENTES, COLUBRIDAE) Albert Schwartz 131 ®(DlL®(g'^ ^mW) IB®^IiSf^ Volume 16, Number 1 December 16, 1969 i.:us. coy.p. zocl. LIBRARY JAMS 1970 VARIATION AND RELATIONSHIP OF THE STUDll^fSHES^RD FUNDULUS CATENATUS AND FUNDULUS ^T^fcfi^^RSlTYj (CYPRINODONTIDAE, PISCES) JAMIE E. THOMERSON Faculty of Biological Science, Southern Illinois University, Edwardsville, Illinois 62025 p. 1 SWINGLEUS POLY cur HROIDES GEN. ET SP. N. (MONOGENEA: GYRODACTYLIDAE) FROM FUNDULUS GRANDIS BAIRD AND GIRARD WILMER A. ROGERS Fisheries Laboratory, Agricultural Experiment Station, Auburn University, Auburn, Alabama 36830 p. 22 A NEW TREMATODE, LEPIDODIDYMOCYSTIS IRWINI N. G., N. SP. (DIDYMOZOIDAE) FROM A MARINE FISH, MENTICIRRHUS NASUS SATYU YAMAGUTI AND SHUNYA KAMEGAI Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana 70118 p. 26 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in chapters I and II (only) of the Style Manual for Biological Journals, 2nd ed., published in 1964 by the American Institute of Biological Sciences, Washington, D. C. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully correaed. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. Illustrations should be proportioned for one or two column width reproductions and should allow for insertion of legend if occupying a whole page. An abstraa not exceeding three percent of the length of the original article must ac- company each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. The editors also recognize the policy adopted by the Federal Council for Science and Technology, and endorsed by the Conference of Biological Editors, that page charges for publication of scientific research results in scientific journals will be budgeted for and paid as a necessary part of research costs under Federal grants and contracts. Accord- ingly, writers crediting research grant support in their contributions will be requested to defray publication costs if allowable under the terms of their specific awards. Illustrations and tabular matter in excess of 20 percent of the total number of pages may be charged to the author; this charge is subject to negotiation. Exchanges are invited from institutions publishing comparable series but subscriptions are available if no exchange agreement can be effected. Separate numbers or volumes can be purchased by individuals and subscriptions are accepted. Remittance should accompany orders from individuals. Authors may obtain separates of their articles at cost. Address all communications concerning manuscripts and editorial matters to the editor; communications concerning exchanges, and orders for individual numbers to Dr. Royal D. Suttkus, Department of Biology, Tulane University, New Orleans, Louisiana 70118. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. and Bot. Price per number (flat rate) : $1.50 Price per volume (flat rate) : $5.00 Meade Natural History Library Tulane University Riverside Research Laboratories Route 1, Box 46-B Belle Chasse, Louisiana 70037 Gerald E. Gunning, Editor Alfred E. Smalley, Associate Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Vickie L. Haddenhorst Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 16, Number 1 December 16, 1969 VARIATION AND RELATIONSHIP OF THE STUDFISHES, FUNDULUS CATENATUS AND FUNDULUS STELLIFER (CYPRINODONTIDAE, PISCES; JAMIE E. THOMERSON Faculty of Biological Science, Southern Illinois University Edwardsville, Illinois 62025 Abstract Fundulus catenatus and Fundulus stel- lifer are recognized as valid allopatric (with the exception of a problematic syntopic record) species differing in pigmentation, male breeding coloration, distribution of male contact organs and pharyngeal denti- tion. On the basis of meristic and mor- phometric differences eight populations (Ozarks, Ouachita, Homochitto River, Ten- nessee R., Virginia, Cumberland R., Green R. and Indiana) of F. catenatus and six populations (LIpper Coosa R., Middle Coosa R., Alabama R., Cahaba R., Tallapoosa R. and Chattahoochee R. ) of F. stellifer are recognized. The Homochitto population seems to be dwarfed and no black-banded males were recorded. In both species, males occur in two breeding color forms: those with and those without a black band on the caudal fin. Fundulus sicllifer males with the black caudal band have black terminal dorsal bands also. The black caudal band is usually subterminal in F. catenatus and terminal in F. stellifer males. The more massive pharyngeal bones and teeth of F. stellifer are regarded as evi- dence for further specialization for snail feeding. Fundulus catenatus probably orig- inated in the Eastern Highlands and the Ozarks and Ouachita populations may be the result of separate invasions of the In- terior Highlands. Fundtdus stellifer was derived from a F. catenatus-like ancestor isolated in the Alabama River system by minor stream capture. Introduction The northern studfish, Fundulus catenatus (Storer) 1846, is a large active killifish which inhabits the clear sand and gravel bottom streams draining the upland areas of the cen- tral United States. It is known only from streams ultimately tributary to the Mississippi River and is one of a number of animal spe- cies distributed disjunctly in the Interior Highlands of Missouri and Arkansas west of the Mississippi River and in the Cumber- land and Appalachian uplands east of the Mississippi River (see Conant, I960, for a discussion of this distributional pattern). With the exception of an isolated population in the Homochitto River drainage in Missis- sippi, F. catenatus is absent from the inter- vening Mississippi Embayment of the Gulf Coastal Plain to the south and the predomi- nantly prairie state of Illinois to the north. Fundulus stellifer (Jordan) 1876, the south- ern studfish, has its main distribution in the upland tributaries of the Alabama River in Alabama and Georgia and few of the head- water tributaries of the Chattahoochee River in northern Georgia. With the exception of a single problematic syntopic record dis- cussed below, the two studfishes appear to be completely allopatric. Though extensive comparative life history data are nor yet available, they occupy similar habitats and F. stellifer may be thought of as replacing F. catenatus in its range. The studfishes are quite similar morpho- logically ( Figure 1 ) and were even con- sidered conspecific by Garman (1895, pp. 107-108), but presently (Moore, 1968; Editorial Committee for this Paper: Dr. Robert R. Miller, Curator of Fishes, University of Michigan, Ann Arbor, Michigan Dr. Stanley H. Weitzman, Curator, Division of Fishes, U. S. National Museum, Washington, D. C. 1 Tulane Studies in Zoology and Botany Vol. 16 Figure 1. A. Male Futidiihis catenatus (SL 86.6 mm), and B. female F. catenatiis (SL 69.4 mm) from SIUE collection JET 66-10, Ozarks Population. C. Male Fundulus .stcUifer (SL 68.5 mm), and D. female F. siellifcr (SL 68.8 mm) from TU 37703, Cahaba River Population. Bailey, et ai, I960) they are considered dis- patric species. The purpose of this study is tinct species. However, Miller (1955) has to describe and compare variation in the two reasonably suggested that further study might forms and to clarify their relationship. The show them to be subspecies rather than alio- problem is simplified in that the studfishes No. 1 Variation and Relationship of Two Stud fishes 97 95 93 91 FUNDULUS CATENATUS • 89 87 86 83 81 BOTH D FUNDULUS STELLIFER O Figure 2. Distribution of studfish collections listed in Material Examined and peripheral lo- calities in Kansas and Oklahoma. have not been described several times, con- fused, or misidentified as has so often hap- pened to members of the genus Fundulus. Even though the two forms show consider- able overlap in meristic and morphometric characters, consistent differences in pigmen- tation, male breeding colors, and pharyngeal dentition lead me to regard F. catenatus and F. stellifer as valid allopatric species. Key to the Studfishes Eight to ten continuous rows of lateral dots centered on each scale; dots brown or olive in females and juveniles, orange to orange-red in nuptial males; scale outlines not sharp, particularly in males; males with- out black margin on dorsal fin but some males with black subterminal caudal band; pharyngeal teeth of large individuals at most peg-like; nuptial males usually without con- tact organs on dorsal fin or caudal peduncle; lateral scale rows usually 12-14. In streams ultimately tributary to the Mississippi River _ Fiindaliis catenatus. Dots randomly distributed or at most in six to nine irregular rows on upper sides, dots not centered on each scale, size of dots irregular; dots brown or olive in females and juveniles, orange to dark red in nuptial males; scale outlines sharp, particularly in Tulane Studies in Zoology and Botany Vol. 16 females and juveniles; some males with black margins on both dorsal and caudal fins; some pharyngeal teeth in large individuals molari- form; males often with contact organs on dorsal fin and caudal peduncle; lateral scale rows usually 14-16. In streams ultimately tributary to the Alabama or Chattahoochee Rivers Fundulus stellifer. Range Distribution of Material Examined and Peripheral Records ( Figure 2 ) gives a good representation of the range of the studfishes. A few of the apparent distributional gaps are not real but reflect that no material from those areas was studied. Fundidus catenates Moore (1968, p. 112) gave the range of Vunduhis catenatiis as: "Upland tributaries of the Tennessee, Cumberland, and Green Rivers in Ken- tucky, Tennessee, Virginia, and Ala- bama: clear streams of the Ozark region in Kansas, Missouri, Oklahoma, and Ar- kansas; Red River tributaries in south- western Arkansas; the Homochitto River in Mississippi; and the upper part of the East Fork of White River, Indiana." Previous records of F. catenatus in Kansas are probably not valid (Cross, 1967, pp. 230- 231), however, Bass and Triplett (1967) have taken a single specimen from Drywood Cr., Crawford County Kansas, a stream ulti- mately tributary to the Little Osage River. The populations cited in this paper represent geographical areas from which samples seemed to me to be reasonable homogeneous and statistically distinct from those of adja- cent areas. The "Ozarks Population" inhabits the small creeks draining into the Mississippi River on the Missouri ( western ) side of the river from just north of the confluence of the Missouri and Mississippi Rivers south to Cape Girardeau Co., Missouri. Smith ( 1965 ) included F. catenatus as a hypotheti- cal addition to the fish fauna of Illinois be- cause of its occurrence in these small tribu- taries in adjacent Missouri counties. The populations north of the Missouri River in the Big Creek system seem to have crossed the river from the south and established themselves in historical times (W. L. Pflieger, in litt., 1966). It thus seems possible that pioneers from the Ozarks population in Mis- souri may someday cross the Mississippi River into Illinois. The specimens included in the Ozarks population are generally distributed throughout the Missouri Ozarks and south into central Arkansas where samples from tributaries to the Arkansas River were re- ferred to this population. I have not exam- ined material from the Neosho R. in NE Oklahoma (Branson, 1967). The "Ouachita Population" includes sam- ples from the Ouachita River drainage in Arkansas and one sample from the Saline River of southwestern Arkansas, a tributary of the Red River. The "Homochitto River Population" is confined to the Homochitto River system in the southwestern corner of Mississippi. The "Tennessee River Population" in- cludes samples from the Tennessee and Duck Rivers and their tributaries in Tennessee, Alabama, and Georgia. CU 50009 (22 speci- mens) from Powell River, Claiborne Co., Tenn., just south of the western tip of Vir- ginia ( see Figure 2 ) clearly belongs with the Tennessee River population and not with the Virginia population. The "Virginia Population" includes sam- ples from the Clinch River and the North Fork of the Holston River in Virginia. Al- though several of the collections I examined had locality data simply as "Holston River, Va." or just as "Va.," I assume that they came from the North Fork of the Holston River or the Clinch River. Ross and Carico (1963) listed 25 localities for F. catenatus from the North Fork drainage and none for the Mid- dle Fork or South Fork of the Holston River. The "Cumberland River Population" in- cludes samples from the Cumberland River and its tributaries in Tennessee and Ken- tucky. The "Green River Population" in- cludes samples from Green River drainage in Kentucky and Tennessee. The "Indiana Population" is a relict population in tribu- taries to the East Fork of the White River in Indiana (see Gerking, 1945, p. 22; Map 69, p. 78). Fundulus stellifer Fundulus stellifer occurs in Alabama River tributaries in Alabama, Georgia and Polk No. 1 Variation and Relationship of Two Stt/dfishes Co., Tennessee, but is not known from the Tombigee River wliich joins the Alabama River from the west just north of Mobile Bay. It also occurs in several of the upper tributaries of the Chattahoochee River in northern Georgia ( Figure 2 ) . There is a problematic record of one specimen from West Fork Chickamauga Creek, a Tennessee River tributary in northeastern Georgia ( see Material Examined). This record is of con- siderable interest because it is from the range of F. catenatus and an F. catenates was pres- ent in the collection. I believe this is a valid record of F. stellijer syntopic with F. cate- natus rather than the result of mixing of specimens from two different localities or some similar mistake. However, intensive collecting of this locality 17 September 1967 netted a single juvenile F. catenatus. All other easily accessible sites in the small drain- age were visited also; two offered no stud- fish habitat, one seemed suitable but no studfish were present and F. catenatus only was present at two other sites. One party of fishermen was present at the syntopic locality when we arrived at 0900 and another party came before 1030. Perhaps the record of F. stellifer at this locality indicates bait release rather than a natural population, but it seems doubtful that fishermen would transport bait from the sparsely populated Alabama River tributaries of the area when bait fishes are so abundant in the Tennessee River tribu- taries. The "Upper Coosa River Population" in- cludes samples from northern Georgia, ex- cept for those from Chattahoochee tributaries. Included were samples from Walker, Mur- ray, Whitfield, Cobb, Gordon and Pickens Cos. The "Middle Coosa River Population" comprises all samples from the Coosa River and its tributaries in Alabama and in Chat- tooga and Cherokee Cos., Ga. The "Alabama River Population" included samples from the Alabama River proper and its small tributaries. The "Cahaba River Population" and the "Tallapoosa River Popu- lations" included samples from tributaries to those streams respectively in Alabama and Georgia. The "Chattahoochee River Popula- tion" includes all samples from the Chatta- hoochee system. Methods All rays of the dorsal, anal and left pec- toral fins were counted at their bases. Caudal ray number is total branched caudal rays plus 2 and counts were made only on large indi- viduals since the number of branched rays increases with ontogeny. Lateral scales were counted as described by Hubbs and Lagler ( 1947). Lateral scale rows were counted on the left side only in a zig-zag fashion from and including the scale on the dorsal midline at the anterior of the dorsal fin base vcn- trally to the last scale before the ventral mid- line. Standard Length measurements were made to the nearest mm for the fish discussed in the section on contact organs. All other mea- surements, including Standard Length, were made with Helios mm dial calipers to three figures. Standard Length (SL), Predorsal Length (PDL), Caudal Peduncle Depth (CPD), Pelvic Fin Length (P2L), Depressed Dorsal Fin Length (DL), Depressed Anal Fin Length (AL), and Anal Fin Base Length ( AB ) were taken as described by Hubbs and Lagler (1947). Body Depth ( BD ) and Head Width (HW) were taken as de- scribed by Thomerson (1966). Head Length (HL) was taken from the rear of the bony operculum at the dorsal terminus of the gill slit to the tip of the snout. Head Depth ( HD ) was taken at the level of the posterior margin of the preopercle. Color descriptions were made from 35 mm color slides of live and preserved speci- mens, from field notes and from live speci- mens kept in aquaria. All adult males in the collections listed under Material Examined were examined with a dissecting microscope and sketches were made showing distribu- tion of contact organs. Notes on pigmenta- tion were also made for these males. Variation in Meristic Characters Meristic data are presented in tabular form (Tables 1-6). Populations are not con- sidered to have significant differences for a given count unless the sum of 2 standard errors (2SE) of one plus 2SE of the other is less than the difference between the two means. This interpretation and that of Fig- ures 5 and 6 presenting morphometric data follow Hubbs and Hubbs (1953). 6 Ttdane Studies in Zoology and Botany Vol. 16 Table 1. Number of dorsal rays in Fundiilus catenatus and Fimduhis stellifer. POPULATION 12 13 14 15 16 17 N Average SD 2SE Funduhis catenatus Ozarks 14 188 58 9 269 15.2 .58 .07 Ouachita 1 5 64 29 1 100 14.2 .60 .12 Homochitto R. 10 71 19 100 14.1 .53 .11 Tennessee R. 13 65 48 2 128 14.3 .54 .09 Virginia 18 38 9 65 13.9 .63 .16 Cumberland R. 3 29 56 12 100 15.2 .81 .16 Green R. 3 45 62 7 117 14.6 .64 .12 Indiana 3 3 6 Fiindulus stellifer Upper Coosa R. 6 30 19 3 58 14.3 .74 .19 Middle Coosa R. 43 76 6 1 126 13.7 .58 .10 Alabama R. 1 27 44 1 73 13.6 .54 .12 Cahaba R. 33 55 1 89 13.6 .50 .10 Tallapoosa R. 1 7 1 9 13.0 Chattahoochee R. 5 14 11 1 31 14.3 .77 .28 Ftmdidus catenatus Observed range of variation in dorsal ray number is 12 to 17 (Table 1). Specimens from the Ozarks and Cumberland River pop- ulations tend to have relatively high dorsal ray numbers (avg. 15.2 for both) and these populations are significantly different from all other populations of F. catenatus in this character. The two populations with lowest average number of dorsal rays are the Homo- chitto River (14.1) and Virginia (13.9) populations. The Homochitto River popula- tion is not separable from either the Ouachita or Tennessee River populations with this character; however, the Virginia and Ten- nessee River populations can be separated. Anal ray number (Table 2) varies from 13 to 18. The highest averages are again those of the Ozarks and Cumberland River populations (avg. 16.2 for both), but the Green River Population also tends toward high counts (avg. 16.1). The Homochitto River, Ouachitas, and Virginia populations tend toward low counts (avgs. 14.9, 15.2, and 15.3 respectively). Distribution of anal ray counts tends to parallel distribution of Table 2. Number of anal rays in Fundulus catenatus and Fundulus stellifer. POPULATION 13 14 15 16 17 18 N Average SD 2SE Fundulus catenatus Ozarks 12 138 48 6 204 16.2 .60 .08 Ouachita 2 10 53 33 2 100 15.2 .72 .14 Homochitto R. 1 13 78 8 100 14.9 .50 .10 Tennessee R. 2 47 68 19 136 15.8 .70 .12 Virginia 2 39 22 1 64 15.3 .57 .14 Cumberland R. 14 59 24 3 100 16.2 .68 .14 Green R. 12 69 19 100 16.1 .55 .11 Indiana 2 4 6 Fundidus stellifer Upper Coosa R. 1 15 34 9 59 14.9 .69 .18 Middle Coosa R. 3 58 59 5 1 126 14.5 .65 .12 Alabama R. 1 31 37 3 72 14.5 .61 .14 Cahaba R. 2 56 30 88 14.3 .52 .11 Tallapoosa R. 4 5 9 14.6 Chattahoochee R. 1 8 21 1 31 14.7 .59 .21 No. 1 Variation and Relationship of Two Studfisbes Table 3. Number of left pectoral ra>'s in Fuiuhilus cateuatti.s aiul Fuudulm stellifer. POPULATION 14 15 16 17 18 19 N Average SD 2SE Fiindtthis catcnafiis Ozarks 38 131 31 200 17.0 .58 .08 Ouachita 25 67 8 100 16.8 .55 .11 Homochitto R. 1 4 21 63 11 100 17.8 .72 .14 Tennessee R. 1 4 41 66 9 1 122 16.7 .73 .13 Virginia 11 44 8 63 16.0 .55 .14 Cumberland R. 1 1 34 61 3 100 16.6 .61 .12 Green R. 7 48 39 6 100 16.4 .71 .14 Indiana 4 2 6 Funduhis stellifer Upper Coosa R. 1 11 38 8 58 16.9 .63 .17 Middle Coosa R. 4 49 60 12 1 126 16.7 .74 .13 Alabama R. 33 39 1 73 16.6 .57 .13 Cahaba R. 8 60 22 90 16.2 .56 .12 Tallapoosa R. 1 1 4 3 9 17.0 Chattahoochee R. 1 13 17 31 16.5 .57 .20 dorsal ray counts; most specimens have one more anal than dorsal ray but some have the same number, one less, or two more. Number of left pectoral rays ( Table 3 ) varies from 14 to 19. The Virginia sample average (l6.0) is significantly the lowest and the average ( 17.8) for the Homochitto River sample is significantly the highest. Co- efficient of Difference between these two populations is 1.38, slightly above the con- ventional criterion for subspecific differen- tiation (Mayr, Linsley, and Usinger, 1953, p. 146). However, Coefficients of Differ- ence between the Homochitto population and the Ozarks, Ouachita, or Tennessee River populations are all less than 1.0. The same is true of Coefficients of Difference between the Virginia population and the Tennessee River, Cumberland River, or Green River populations. Caudal Ray number (Table 4) varies from 15 to 18. All populations tend to have sub- equal proportions of individuals with 16 or 17 caudal rays. The lowest average number of caudal rays is 16.2 (Green River) and the highest is 16.7 (Homochitto River). There is thus a weak trend from higher num- ber of caudal rays in the south to lower num- ber of caudal rays in the north. Observed range of lateral scale row num- Table 4. Number of caudal rays in Funduhis catenatu.s and Fundulm stellifer. POPULATION 14 15 16 17 18 N Average SD 2SE Fundidus catenatus Ozarks 2 57 24 3 86 16.3 .58 .13 Ouachita 3 17 18 3 41 16.5 .74 .23 Homochitto R. 18 22 4 44 16.7 .63 .20 Tennessee R. 1 19 24 2 46 16.6 .61 .18 Virginia 11 7 18 16.4 .25 .12 Cumberland R. 3 20 27 1 51 16.5 .64 .18 Creen R. 6 31 13 2 52 16.2 .69 .19 Indiana 6 6 Funduhis stellifer Upper Coosa R. 1 11 17 29 16.6 .57 .21 Middle Coosa R. 5 26 27 58 16.4 .64 .17 Alabama R. 1 36 17 54 16.3 .56 .15 Cahaba R. 1 17 33 1 52 15.7 .58 .16 Tallapoosa R. 6 3 9 16.3 Chattahoochee R. 5 14 4 23 16.0 .64 .27 8 Tulane Studies in Zoology and Botany Vol. 16 OZARKS 5 OUACH 5 3 HOMO- CHITTO 4 2 3 TENN. R. 5 4 4 3 VA. 3 3 4 4 4 CUMB. R. 4 4 5 5 4 1 GREEN R. 4.3 3.5 4.0 3.5 4.0 3.2 3.8 X Figure 3. Matrix giving the number (out of 6) of meristic characters in which the various populations of Ftmdulus catenatus differ sig- nificantly. X is the arithmetic average number of significantly different characters for each population. Population labels apply to the hor- izontal row to the left of the lal3el and the ver- tical column below the label. ber (Table 5) is 11 to 16. Average number of lateral scale rows ranges from 13.2 (Ouach- ita, Green River) to 14.4 (Virginia). The Virginia sample average differs significantly from the Tennessee River average (13.7). Number of lateral scales is given in Table 6. In the aquarium studfish fight and indi- viduals may be almost completely scaled. Though such intensive fighting may be an artifact caused by crowding in the aquarium, many studfish collected in the wild show patches of regenerated scales. These indi- viduals were not counted unless the original number of scales was obvious. However, there is also considerable irregularity of scale rows that does not involve regenerated scales. When I felt that the lateral scale count was questionable, I counted the lateral scales on the right side and discarded the counts if there was more than 1 scale difference. Thus, even though the count is often difficult to make, the wide range of variation (38-52) seems real. The average (42.4) for the Homochitto River Population is significantly the lowest. Other populations with low aver- age number of lateral scales are the Ozarks (avg. 43.5) and Ouachita (avg. 44.5) pop- ulations. These southern and western popu- lations thus contrast with the eastern popu- lations which have averages ranging from 45.3 to 46.4. These differences are, however, far below the level required to justify recog- nition of subspecies. Figure 3 is a matrix showing the number of meristic characters (out of six) in which the various populations of F. catenatus differ. The average number given for each popula- tion is the arithmetic mean and is an un- weighted estimate of the amount of diver- gence of each population from all other F. catenatus populations. The Ozarks popula- tion is most divergent and the Cumberland River population the least divergent. Table 5. Number of lateral scale rows in Fundtthis catenatus and Funduhis stelUfer. POPULATION 11 12 13 14 15 16 17 18 N Average SD 2SE Fundulus catenatus Ozarks 6 70 112 12 200 13.6 .68 .10 Ouachita 20 46 31 3 ino 13.2 .78 .16 Homochitto R. 1 22 29 37 10 1 100 13.4 1.00 .20 Tennessee R. 6 35 67 13 1 122 13.7 .74 .13 Virginia 8 24 23 5 60 14.4 .99 .26 Cumberland R. 2 12 42 37 7 100 13.3 .85 .17 Green R. 15 48 37 100 13.2 .69 .14 Indiana 2 4 6 Fundulus steUifer Upper Coosa R. 1 11 21 13 2 48 15.1 .87 .25 Middle Coosa R. 3 27 36 35 17 1 119 15.4 1.09 .20 Alabama R. 12 46 12 3 73 15.1 .70 .16 Cahaba R. 5 42 26 14 87 15.6 .83 .18 Tallapoosa R. 6 2 1 9 15.4 Chattahoochee R. 1 7 15 6 2 31 15.0 .91 .33 No. 1 Variation and Relationship of Two Stud fishes S 3 !*< 15 C 03 CO 3 ■+^ C c a 3 s 3 o a s z CO a M CS > < IC CO CO o o CD in CO 00 -H CO o O CD CM CO CO CO >— I 1 — I ^^ CO i-H (M (N in CD (M CO Co 3 iX z 1.^ tf o s o *-• aj H 4-* ,12 * < o ^ o o V3 "5 o CO OS o CJ 3 O S o C c > CL, ^ O) -^ ^ O -f CD CO cq o o CO CO in -f (N c— I 1-H I— I 1—1 in ^ i^ 05 ^ in t^ (M CO ^ (M 00 00 m CD CO (M in ^ U O ►:: CD oc in in 05 o in CO CO I- CO oq (N ^ 1^ (N t^ 00 CO 05 in t- oi oi in CO CD CO ^^ CO in ^ "* CD t^ CO cni i> 00 CM CO in -t CO CM ^ m CD -* in CM CM —I 1— I >-H in -* cs in CM ^ CO t^ ^ CM CO CO .-H CO tf ^ 15 -2 4-' <../ "cs "« rt -C s < U H U 10 Tulane Studies in Zoology and Botany Vol. 16 u COOSA \ 2 M COOSA k 2 1 ALA. R. 6 2 3 CAHABA 2 2 2 4 CHAT'E 3.0 1.8 2.0 3.8 2.5 X Figure 4. Matrix giving the number ( out of 6) of meristic characters in which the various populations of Ftinduhis steUifer differ signifi- cantly. Interpretation as for Figure 3. Fundulus SteUifer Fundulus SteUifer populations from the Alabama River-Coosa River system show a trend toward high average number of rays in all fins (pelvic not counted) upstream and low average number of rays downstream. The Cahaba River sample tends to have the lowest average numbers of fin rays. Number of dorsal rays ( Table 1 ) varies from 12 to 16 and all populations have a modal number of 14. However, means ( 14.3 ) for the Upper Coosa River and Chattahoochee River samples are signifi- cantly higher than means for the other popu- lations. Number of left pectoral rays ( Table 3) varies from 15 to 19 and the Cahaba River average (16.2) is significantly lower than averages for other Alabama drainage populations. Caudal ray number ( Table 4 ) varies from 14 to 17. With the exception of the Cahaba River and the Chattahoochee River samples, individuals with 16 or 17 caudal rays occur in all populations in sub- equal proportions. These samples have un- usually high percentages of individuals with 15 caudal rays and the Cahaba River average (15.7) is significantly the lowest for the Alabama drainage samples. Trends in distribution of average numbers of scales do not parallel distribution of fin ray averages. Left lateral scale row number varies from 13 to 18 (Table 5). The Cahaba River sample has the highest average num- ber (15.6) and the Chattahoochee River sample the lowest (15.0). The remarks above regarding the causes of extreme variation in lateral scale number of F. catenatus also apply to F. SteUifer. Observed range of variation in lateral scale number (Table 6) is 38 to 52. The Cahaba and Tallapoosa River sam- ples have the lowest average number of lat- eral scales (42.7, 42.2) and the Middle Coosa River sample the highest ( avg. 45.9). Figure 4 is a matrix for F. steUifer drawn in the same fashion as Figure 3. The Cahaba River sample has the highest average num- ber of significantly different meristic char- acters and the Middle Coosa River sample the lowest. Comparison of Fundulus catenatus and Fundulus SteUifer Dorsal Rays — Fundulus steUifer tends to have fewer dorsal rays than F. catenatus but there is almost complete overlap in observed range of variation (none with 17 rays for F. SteUifer, 9 from the Ozarks sample for F. catenatus). The Middle Coosa River, Ala- bama River, Cahaba River and Tallapoosa River samples of F . steUifer have lower aver- age numbers of dorsal rays than any F. cate- natus sample (Table 1), but the Upper Coosa River and Chattahoochee River samples have higher average numbers than three of the F. catenatus samples and are identical to the Tennessee River sample (avg. 14.3) in this parameter. Anal Rays — The Homochitto River F. catenatus sample is not separable from the Upper Coosa River and Chattahoochee River F. SteUifer samples of this character. The other three large samples of F. steUifer had lower averages than all F. catenatus and the other six large F. catenatus samples had higher averages than all F. steUifer samples. Left Pectoral Rays — Range of variation in average number of left pectoral rays (16.0 to 17.8) for the F. catenatus samples includes the range of variation (16.2 to 16.9) seen in F. SteUifer samples. Caudal Rays — Only the Cahaba River F. SteUifer sample average (15.7) is signifi- cantly outside the range of variation in this parameter for F. catenatus samples ( 16.2 to 16.7). No. 1 Variation and Relationship of Tti'o St/ulfishes 11 PDL/SL BD/SL CPD/SL fUWDUlUS CATtNATUS SEX I M O Z A f)K S f HOMOCHITTO H TE NNE SSE ( R CUMBERLAND R fUNOULUS STELLIFtR U COOSA R M COOSA R ALABAMA R CAHABA n CHATTAHOOCHEE R ^ ^JZ ^^ T^ 630 650 670 690 170 190 210 110 120 130 140 Figure 5. Geographic variation in morphometric characters in Fiindiilus catenatus and Fiin- diilus stellifcr. Samples are 10 indix'iduals except where nunilDcr is given. PDL-Predorsal Length, BD-Bodv Depth, CPD-Candal Peduncle Depth, SL-Standard Length. Drawn following Hubbs and Hubbs (1953). Lateral Scale Rows — There is considerable overlap in range of variation in this charac- ter (F. catenatus, 11 to 16; F. stellifer, 13 to 18), but it is the most divergent of the meristic characters studied. The highest average for an F. catenatus sample ( 14.4, Virginia Population ) is significantly lower than the lowest F. stellifer average (15.0, Chattahoochee River Population ) . Lateral Scales — The range of average num- ber of lateral scales for F. catenatus samples (42.4 to 46.4) includes the range of varia- tion (42.7 to 45.9) for F. stellifer samples. Morphometric Characters There is considerable variation in body proportions in both species of studfish. Sev- eral characters also exhibit marked sexual dimorphism. In addition to the characters shown in Figures 5, 6, and 7, other propor- tional measurements involving characters mentioned above in the section on Material and Methods were made but showed only random variation and are not presented. There is striking sexual dimorphism in Predorsal Length ( Figure 5 ) in F. catenatus and to a lesser extent in F. stellifer. Even though the body outline of males and females is different ( Figure 1 ) there is no sexual dimorphism of Body Depth in either species but males do tend to have deeper caudal peduncles than females. This character is fairly consistent from population to popula- tion in F. catenatus, but there is considerable variation in F. stellifer. Male F. stellifer from the Coosa River and Alabama River popula- tions tend to have deeper caudal peduncles than most F. catenatus males. The Cahaba River population resembles F. catenatus in this character and the Chattahoochee River population appears most slender. Head Length ( Figure 6 ) varies consider- ably within populations but there does not seem to be a consistent pattern of either sexual or geographic variation in F. catenatus except that Tennessee River females have significantly shorter heads than males. The Alabama River F. stellifer also show signifi- 12 Tulane Studies in Zoology and Botany Vol. 16 OL/SL MOUHJS C ATtNATUS StK M 01 ARKS r UA CHITA5 MOMOCMITTO I TINNESSE I CUMB i R LAND R fUNOULUS STULIFtW U COOSA R CAHABA R CHAT TAHOOCHIE R T ^^ .340 .380 t; i^ .500 .180 .220 260 300 .340 .380 .220 .240 .260 .280 .300 Figure 6. Geographic variation in morphometric characters in Fiinduhis catenatus and Fun- dulus stellifer. Samples are 10 individuals except where number is given. HL-Head Length, BL- Body Length, DL-Depressed Dorsal Fin Length, AL-Depressed Anal Fin Length, SL-Standard Length. Drawn following Hubbs and Hubbs ( 1953). cant sexual dimorphism in this character and difference between the Alabama and Cahaba River populations is significant when males are compared with males and females with females. There is also a significant difference between Cahaba and Chattahoochee River populations. Dorsal Fin Length ( Figure 6 ) shows a high degree of sexual dimorphism with little overlap between the sexes in either species. With the exception of Ozarks population F. catenates, there is little variation in female dorsal fin length. Part of the variation seen in the males is seasonal. Nuptial males have slightly elongated dorsal fins. Male studfish have dorsal and anal fins of approximately the same length but anal fin length averages 17% greater than dorsal fin length in female F. catenatus and 13% greater in female F. stellifer. The anal fin of the female digs into the substrate during spawn- ing and her anal rays are stouter than the dorsal rays. In female studfish longer anal fins tend to be elongate and pointed but shorter anal fins tend to be rectangular. The Tennessee River females have significantly shorter anal fins than those of other F. cate- natus populations. The Homochitto River females have significantly longer anal fins than females from other eastern populations of F. catenatus. The means for F. stellifer samples, except the Middle Coosa River sam- ple, are significantly lower than those of F. catenatus samples except for the Tennessee River population. None of the F. stellifer samples can be separated from the Tennessee River F. catenatus sample using this charac- ter. The Cahaba River sample mean is sig- nificantly lower than the other F. stellifer sample means except for the Upper Coosa River sample mean. Both species show sexual dimorphism in Pelvic Fin Length. The observed range of P2L/SL for F. catenatus females is 0.102 to 0.136, for F. stellifer females 0.106 to 0.131, average for both 0.115. This average ratio is compared with the distribution of P2L/SL for males in Figure 7. Large males and No. 1 Variation and Relationship of Two Stud fishes 13 16 15. 14. Z 13 c « -I 12J 11. — 10. « a. 7. 6. 40 O o o o. * .o8°o *• ^. o° .°:-- «Qi4 • • O O 1^ 60 70 80 90 Standard Length in mn i6o 120 Figure 7. Relationship between Pelvic Fin Length and Standard Length in Fiitidiihis cate- natus and Fundulus stellifer. Pooled data from all populations, including largest males from each. smaller males in breeding condition tend to have proportionately longer pelvic fins in both species. Male Contact Organs and Breeding Colors During the breeding season, males of F. catenati/s and F. stellifer develop contact organs (Newman, 1907, 1909). These are particularly noticeable on the anal fin rays, and the presence of contact organs on the anal fin of F. stellifer was noted by Fowler (1916, PI. 1). The distribution of contact organs is similar in the two studfishes ( Fig- ure 8 ) , but usually F. catenatus males are less "prickly" than F. stellifer males, and contact organs on the sides of the body are confined to an area between the bases of the dorsal and anal fins. Figure 8A shows the distribu- tion of contact organs usually seen on breed- ing F. catenatus males, but there are often no contact organs on the dorsal or pectoral fins in this species. In both species a few anal fin contact organs may be present in males taken at any time of year. As the fish come into breeding condition, the anal fin organs are proliferated and contact organs on the cheeks and opercles develop concurrently or soon after. Development of the lateral con- tact organs begins at midbody between the bases of the dorsal and anal fins and in gen- eral proceeds further anteriad and posteriad 14 Tulane Studies in Zoology and Botany Vol. 16 Figure 8. Sketches of stiidfish males; dotted area represents extent of development of contact organs. A. Usual extent of contact organs in Fundulus caienatus. B. Unusually extensive devel- opment of contact organs in Fundulus catenatus. C. Usual extent of contact organs in Fundulus stellifer. in F. stellifer than in F. catenatus ( Figure 8A and C ) . I have seen only two specimens of F. catenatus with contact organs on the lateral scales of the caudal peduncle; one of these is sketched in Figure 8B. Development of contact organs on the pelvic rays, on both sides of the fin, is seen in most high males but presence of contact organs on the pec- toral fins is less common. About half the breeding males of both species have a few organs on the anteroventral surface of the pectoral fin. Occasional males have one or two contact organs on the posterodorsal sur- face of the pectoral fin. Less than half the F. catenatus males examined had contact organs on the dorsal fin, and these were generally confined to the anterior half of the fin. Almost all F. stellifer males had dorsal fin contact organs with the greatest concen- tration in the posterior half of the fin. I examined a number of breeding female stud- fish and found no contact organs. Female studfish do not show seasonal color changes. The patterns shown in Figure IB and D are seen in live specimens in varying shades and intensities of brown. Nonbreed- ing males resemble the females in drab color- ation and pattern, but nuptial males are spectacularly colored. Fundulus catenatus males occur in two color forms. As given in Table 7, various percentages of males exhibit a subterminal black band in the caudal fin. This band in- tensifies during the breeding season but is discernable in all months of the year. Place- ment and sharpness of the band are variable. Some males, particularly those from the Cumberland and Green rivers, have a wide ( about 2 mm ) yellow to orange band distal to the black band. In others, particularly those from the Ouachita and Ozarks popu- POPULATION Number Percent with Band Fundiilus catenatus Ozarks 109 44 Ouachita 12 42 Homochitto 60 0 Tennessee 30 20 Virginia 14 36 Cumberland 30 43 Green 34 12 Ftindtdus steUifer All 142 58 No. 1 Variation and Relationship of Two Studfishes 15 Table 7. Occurrence of males with black reflected light. Males without the black sub- caudal rm band m tunatilus catenatus and ,^^^; i u j u-u- l n Fundidus stcUifcr. terminal band exhibit the yellow to orange terminal band on the caudal fin. The head pattern is similar to that of F. SteUifer. There is a very conspicuous light blue or blue-green iridescent patch from the upper anterior quarter of the orbit to the corner of the gape. The iris is bright yellow- green to dark with a silver reflecting margin around the pupil. In some specimens there is an area of dark pigment directly ventral to the preocular light patch. The cheeks and opercular area are marked with a random pattern of circular orange spots. Those on the operculum are about twice the diameter of the others on head and body. The lower half of the head and the gular and branchi- ostegal regions are yellow, but the upper lations, the yellow to orange band may be lateral surfaces of the head are livid yellow- quite narrow. In two of the Ouachita speci- green. The dorsum and top of the head are mens, the black band is essentially terminal, olive grading into either sky blue or green as it is in F. steUifer. The margins of the laterally. The color of the sides depends on band, particularly the distal margin, are usu- the angle and quality of the light striking ally sharp, but in some males it is a black them. There may be an area of disordered zone rather than a sharply defined band. The orange spots just behind the head or all the band is represented by disconnected blotches spots may be linearly arranged, one spot per of black pigment in many males from the scale, from the gill openings to the bases of Ozarks population. the caudal rays. The spots are slightly sepa- Only one of the males examined showed rated but give the impression of 10 to 12 any concentrated black pigments at the mar- continuous orange lines down the sides, gin of the dorsal fin ( CU 49857, Cumber- Viewed in the water in sunlight against a land River population ) . His black dorsal dark background, a male in high color gives pigment resembles that of an F. steUifer the impression of a fish with a solid red- male with less than maximum expression of orange body. this character. Dorsal fin coloration in breed- Fundulus steUifer males also occur in two ing F. catenatus males varies from dark slate color forms. A black margin on the dorsal blue overall, which appears blue in the wa- and caudal fins may be either present or ab- ter, to yellow distally and translucent yellow- sent. Data from all populations are pooled grey proximally, which looks yellow in the in Table 7, but males of both kinds were water. When the dorsal fin is dark there is present in all populations where reasonable sometimes a narrow yellow or orange margin size samples were available. A few males but I have never seen the white or cream with black borders on the dorsal and caudal margin of the dorsal and caudal fin described fins also had less distinct black borders on by Carman (1895). The interradial mem- the pectoral and pelvic fins, branes of the dorsal and the proximal halves Body color in males is a livid sky blue, of the anal, caudal, and sometimes the pelvic grading into white on the belly and bluish- and pectoral fins are marked with elliptical brown on the dorsum. The fins are clear orange spots. These are largest and most yellow, and there are rows of elliptical orange intense in the dorsal fin. The anal fin is spots in the interradial membranes of the transparent yellow -grey proximally and bright dorsal fin and in the basal halves of the translucent yellow distally. The color pattern caudal and anal fins. There may also be a of the pectoral and pelvic fins is similar but few such spots near the base of the pectoral not so intense. The caudal fin is greyish and fins. If the caudal fin border is absent the translucent in transmitted light but blue in caudal fin may be slate grey rather than yel- 16 Tulane Studies in Zoology and Botany Vol. 16 low. There is a brilliant iridescent pre-ocular patch as in F. catenatiis, but it is yellow-green rather than bluish. The iris is also yellow- green, and there is a light blue-green, diffuse iridescent patch on the gill cover behind the eye. The orange spots on the head are ir- regular in size and arrangement like those of F. catenatiis and the body pattern is similar to the head pattern. Dark orange spots are concentrated on the dorsolateral region of the body with only a few scattered on the lower sides. The spots on the caudal pedun- cle are linearly arranged but not so consist- ently as those of V. catenatiis. Other Characters Male studfish average larger than females in most collections; however, the largest ¥. catenatiis I have seen was a 118 mm SL fe- male from the Ozark population. Dr. David A. Etnier kindly informed me of an unusu- ally large 98 mm SL male F. stellifer from the Conasauga River in Polk Co., Tennessee. The largest males in the collections listed under Material Examined were 86 or 87 mm SL. With the exception of the Homochitto population, a large F. catenates male is over 100 mm SL. The Homochitto population seems dwarfed and the largest specimen (of 768 ) from there was an asexual individual 76 mm SL. Fiindidus catenatus has regular rows of spots on the sides of the body. These dots are red or red-orange in breeding males but an alcoholic male specimen appears plain because of leaching of pigment by the pre- servative. The pattern in F. stellifer is simi- lar but with fewer spots not necessarily on the centers of the exposed scale fields nor arranged in the orderly manner seen in F. catenatus. The few Tallapoosa River speci- mens seem more F. catena! us -\'\ke than speci- mens from other F. stellifer populations in the arrangement of their lateral spots. In addition to the difference in arrangement of the lateral dots there is a consistent differ- ence in the pigmentation of the scale borders. As seen in Figure 1, F. stellifer has a regular cross-hatched appearance in contrast to the more uniform background pattern of F. cate- natus. The cross-hatching extends to the base of the anal fin in F. stellifer and stops a few scales short in F. catenatus. The differences in development of scale border pigmentations al- low identification of juveniles, although they are more similar than are the adults ( Figure 9 ) . I have not seen enough very small speci- mens of the two species to point out differ- ences between them at sizes below approxi- mately 14 mm SL. The intestinal contents of F. stellifer seem finely ground in comparison to those of F. catenatus, probably as a result of chewing with the stout pharyngeal molars ( Figure 10 ) . The pharyngeal bones of F. stellifer are massive compared to those of similar size F. catenatus and molars are developed both on the inferior pharyngeal bones and on the pharyngobranchials. The corresponding teeth in F. catenatus are pegs with hooked tips. They resemble the hooked blade-like teeth of other species of Fundulus, for example F. notatus, but are stronger with an expanded circular cross section. Snails form an impor- tant item of diet for F. stellifer (J. D. Satter- f ield, personal communication ) and I have seen snail shells in the feces of F. catenatus. One might guess from its comparatively more massive pharyngeal bones and denti- tion that of the two, F. stellifer eats more mollusks. Discussion Fundulus catenatus is an aggressive and adaptable small-stream fish widely distributed in the upland areas of the Mississippi Valley. It probably originated in the eastern uplands where I have recognized four distinct popu- lations in roughly a third of the range. The distribution of F. catenatus crosses a number of zoogeographic barriers. Particularly strik- ing is the uniformity of the Ozarks popula- tion over an area approximately equal to the eastern uplands. Many Ozarkian fishes have been sufficiently isolated in the White River system and the Missouri Ozark streams trib- utary to the Missouri and Mississippi Rivers to diverge even to the species level (Gilbert, 1964, pp. 104-5), but F. catenatus samples from the two areas are not significantly dif- ferent. The broadly-ranging Ozark population is the most divergent in terms of meristic char- acters studied ( Figure 3 ) . It seems closest to the Cumberland River population, than to the Tennessee and Green River popula- tions. Not only does relative morphological uniformity over such a large area suggest No. 1 Variation and Relationship of Two Stitdfishes 17 Figure 9. Juvenile studfish— Fi/rirf«/«s catenatus: A. TU 15132 (SL 24.6 mm), B. KU 6661 (SL 24.1 mm), C. KU 11550 (SL 24.2 mm). FuuduJus stellifer: All from JDS, D. SL 22.1 mm, E. SL 25.4 mm, F. SL 17.2 mm. that the Ozarks population is of recent origin compared to the populations in the eastern uplands, but there is evidence that Ozarkian F. catenatus are still rapidly moving into suitable habitats around the margins of the Ozarks. As mentioned above, F. catenatus seems to have crossed the Missouri River and become established in the Big Creek system in recent historical time. The recent Neosho River records may be from a population es- tablished naturally from the White River (Branson, 1967) or from bait releases (Hall, 1956). In any case, F. catenatus is common there now. It seems significant that F. cate- natus is absent from the Flint Hills region of Kansas, even though several other species with Ozarkian affinities occur there (Cross, 1967 ). Probably F. catenatus was not present in the Ozarks when access to the Flint Hills was possible. The Ouachita population shows surprising divergence from the Ozarks population, is most like the Tennessee River population and is not particularly divergent from the other eastern populations. It thus seems probable that the Ozarks and Ouachita popu- lations originated from separate invasions of the Interior Highlands by fish from the east- ern populations. The widely disjunct Homochitto popula- tion does not show as much divergence in meristic characters as the Ozarks population but seems unique in the absence of black- banded males and the small maximum adult size. Conditions in the Homochitto drainage are considerably different from those in the Interior Highlands or eastern uplands and the characters mentioned above could have strong adaptive significance. The small size of the Homochitto fish may be correlated with reduced food supply. In any case, the numbers taken in the collections I have ex- 18 Tulane Studies in Zoology and Botany Vol. 16 PB 2 PB 2 I mm Figure 10. Pharyngeal arches of studfish; A. and B. from a 71.4 mm SL Fundulus stellifer (TU 35029), C. and D. from a 70.0 mm SL Fundulus caienaius ( SIUE, JET 66-10). A. Right lower arch, dorsal view, anterior down. B. Right upper arch, ventral view, anterior to the upper left. C. Left upper arch, ventral view, anterior to upper right. D. Left lower arch, dorsal view, anterior down. amined indicate that F. catenati/s is success- ful in maintaining relatively dense popula- tions in the Homochitto River system. The small fish and southern location of this pop- ulation suggest a similarity to F. stellifer, but their pharyngeal teeth, lateral scale row count and pigment patterns show no such similar- ity. Perhaps gene flow with other popula- tions of F. catenatus is maintained even though there is a broad expanse of unsuit- able habitat separating the Homochitto River from the F. catenatus populations in the cen- No. 1 Variation and Relationship of Two Studfisbes 19 tral highlands. The Homochitto fish may represent a postglacially isolated relict of a population distributed through the Lower Mississippi Valley during glacial periods when lowered sea level caused rejuvenation of small tributaries of the Lower Mississippi River. Alternatively the population could have been established by individuals tribu- tary-hopping during interglacial periods when the Mississippi River was cooler, clearer and swifter flowing than it is today. If this were the case, the Homochitto popu- lation might have received immigrant indi- viduals from both eastern and western populations. The major observed differences between F. catenates and F. stellifer are: general pig- mentation, distribution of contact organs, male breeding colors and pharyngeal denti- tion. F/nid/d/is stellifer could be duplicated by relatively slight changes in an F. catenatiis stock and indeed a few F. catenates individ- uals, as noted above, show tendencies toward the F. stellifer condition. Pharyngeal denti- tion is very similar in small individuals of the two species. Greenwood (1964) found differences of approximately the same mag- nitude between the pharyngeal bones and associated structures of wild individuals and an aquarium-raised specimen of the snail- eating cichlid Astatoreocbromis alluaudi. The aquarium-raised specimen had an adequate diet but did not receive snails until late in life. Its pharyngeal teeth and bones were less massive (Figure 3, p. 5, Greenwood, 1964) than those of any of the wild indi- viduals. The differences between the pharyn- geal teeth and bones of larger F. catenatus and F. stellifer ( Figure 1 0 ) may thus be cor- related with difference in food habits rather than with difference at the genetic level. Whatever their cause, differences between the two species seem to be consistent. The ancestors of F. stellifer were probably a stock much like present day F. catenates which became isolated in the Alabama River system. Whether this was the result of a major faunal exchange between the Tennes- see and Alabama systems or the result of local stream capture is not clear, but the lat- ter seems more likely. If F. catenatus had entered the Alabama system in large num- bers, stabilizing selection acting on the some- what generalized and flexible F. catenatus gene pool would probably have produced just another population of F. catenatiis, one perhaps resembling the Homochitto popula- tion. If the ancestor of F. stellifer was a F. cate- natus-Vike fish, the more massive pharyngeal dentition of F. stellifer is thus probably a result of further specialization for snail eat- ing. It is my impression that, even though the faunas of the Alabama and Chattahoochee systems are rich in fish species, the density of small fishes in streams inhabited by F. catenatus is much higher than in streams in- habited by F. stellifer. The varieties and kinds of habitats seem to be similar in the two areas, so perhaps the greater number of individual fishes in F. catenatus habitat is correlated with availability of food orga- nisms. If F. stellifer lives in a less rich habi- tat, further specialization of snail eating would be of adaptive value by increasing utilization of a food supply little used by other small stream fishes. The evolution of F. stellifer may have been relatively rapid and recent because the characters separating it from F. catenatus seem of obvious adaptive significance but characters of less obvious adaptive significance show little divergence. Material Examined Collections are listed by population, state, county and museum number. Collections de- posited Illinois Natural History Survey (INHS) and Southern Illinois University, Edwardsville ( SIUE ) are not given catalog numbers. Complete data are given for the syntopic collection. Other museum abbrevia- tions are used as follows: The Academy of Natural Sciences of Philadelphia, ANSP; Auburn University, AU; University of Ala- bama Ichthyological Collection, UAIC; Cor- nell University, CU; Indiana University, lU; University of Kansas, Museum of Natural History, KU; University of Michigan, Mu- seum of Zoology, UMMZ; Smithsonian In- stitution, USNM; personal collection of James D. Satterfield, JDS; Tulane Univer- sity, TU; University of Richmond, UR. Number of specimens in each collection is given in parentheses. All specimens were examined and counts were made on 885 F. catenatus and 386 F. stellifer from the col- lections listed below. Distribution of these collections is shown in Figure 2. 20 Tulane Studies in Zoology and Botany Vol. 16 Allopatric Fundulus catenatus Ozarks. — Missouri: Lincoln Co.; SIUE (14), SIUE (44)— Franklin Co.; LXHS (7)— Carter Co.; UMMZ 104684 (25)— Wright Co.; INHS (,3)_Osage Co.; INHS (6)— Phelps Co.; INHS (25)— Morgan Co.; KU 6501 (11)— Camden Co.; UMMZ 150336 (14)— Dallas Co.; KU 7837 (19)— Hickory Co.; KU 7811 (6)— Craw- ford Co.; UMMZ 148448 (9)— Dent Co.; INHS (9), INHS (7), KU 7614 (11)— Wayne Co.; UMMZ 117246 (12)— Cape Girardeau Co.; INHS (8), INHS (2), INHS (25)— Madison Co.; INHS (6)— Shannon Co.; KU 7652 (10), KU 7674 (2), UMMZ 102491 (38)— Dade Co.; KU 5503 (50)— Green Co.; KU 7856 (19) —Taney Co.; KU 10926 ( 112)— McDonald Co.; CU 42324 (6). Arkansas: Sharp Co.; UMMZ 116378 (40)— Marion Co.; KU 8012 (53), KU 9864 (31)— Boone Co.; UMMZ 123522 (88), KU 6661 (8)— Newton Co.; UMMZ 175430 (6)— Madison Co.; UMMZ 123892 (43), UMMZ 123425 (54), UMMZ 123859 (27)— Washington Co.; UMMZ 127769 (7), CU 36506 (2). Ouachita.— Arkansas: Pulaski Co.; INHS (2) — Sahne Co.; CU 42245 (17)— Garland Co.: TU 14250 (3)— Montgomery Co.; KU 6166 (25), KU 6143 (7)— Pike Co.; KU 6133 (48). UMMZ 81112 (8), KU 6135 (11)— Howard Co.: KU 6115 (5)— Polk Co.; KU 6907 (10), UMMZ 81110 (2). Homochitto River. — Mississippi: Amite Co.; UMMZ 144728 (2), UMMZ 155340 (12), TU 7447 (18)— Franklin Co.: TU 11991 (12), TU 16623 (142). TU 19766 (57). TU 19851 (32). TU 15132 (141). TU 23952 (78), UMMZ 161204 (4), TU 7208 ( 176)— Lincoln Co.: TU 32916 (32). TU 28895 (60)— Copiah Co.; UMMZ 154371 (2). Tennessee River. — Tennessee: Sevier Co.; KU 8935 (D— Coffee Co.: TU 30299 (61)— Bedford Co.: UMMZ 121529 ( 1 )— Lincoln Co.: TU 30289 (7)— Marshall Co.; KU 10684 (12) —Marshall and Maurv Cos.; UM\fZ 121477 (44)— Lewis Co.: UMMZ 105213 (28)— Wayne Co.: UMMZ 177703 (4)— Humphrev Co.: UMMZ 168314 (33)— McNairy Co.: TU 10'=;'?3 (2). Tennessee: Claiborne Co.: CU K0009 (22). Alabama: Franklin Co.; UMMZ 177723 (6). Virginia. — Virginia: ? Co.: ( Holstou R.), ANSP 7304-35 (32)— Smvth Co.: UR 1979 (1), UR 899 (2). UR 940 (1), USNM 40368 (3), UMMZ 96899 (11). USNM 102428 ( 1 )— Washington Co.; UMMZ 130779 (1), UMMZ 130808 (4), UMMZ 130788 (3)— Scott Co.: CU 50078 (1), ANSP 74207 (4), ANSP 74235 (D— Lee Co.; USNM 104043 (1). Cumberland River. — Kentucky: Pulaski Co.; CU 50059 (26)— Wayne Co.; CU 50090 (8)— Cumberland Co.; CU 50042 (3), KU 11554 (1), INHS (2). Tennessee: Fentress Co.; KU 11532 (14), CU 50191 (6)— Overton Co.; UMMZ 125196 (2)— Clay Co.; UMMZ 125228 (3), UMMZ 125258 (13), UMMZ 125341 (3) — Jackson Co.; KU 11550 (24), CU 50028 (9)— Smith Co.; CU 49857 (19)— Putnam Co.; KU 8947 (1)— Warren Co.; UMMZ 175257 (5)— Cannon Co.; KU 11678 (10)— Sumner Co.; UMMZ 96323 (26)— Wilson Co.; KU 11588 (15)_Rutherford Co.; KU 11482 (7)— Mont- gomery Co.; INHS (1)— Houston Co.; INHS (8). Green River. — Kentucky: Lincoln Co.; KU 5722 (17), KU 7253 (41)— Casev Co.; UMMZ 178028 (17)— Taylor Co.; KU il620 (26)— Allen Co.; KU 11641 (4), INHS (20), INHS (13). Tennessee: Macon Co.; UMMZ 165377 (42), TU 14532 (25). Indiana.— Indiana: Shelby Co.; lU 397 (5) —Johnson Co.; INHS (2). Syntopic Collection Tennessee River. — Georgia: Walker Co.; W Fork Chickamauga Cr. 1.3 mi. W jet. Hwy. 143 and Hwy. 341 on Hwy. 143, 23 June 1964, R. D. Suttkus and Env. Biol. Class, TU 34975 (1 (5 F. stellifer, 1 9 F. catenatus). Allopatric Fundulus stellifer Upper Coosa River. — Georgia: Pickens Co.; TU 38346 (9), TU 40732 (9)— Murray Co.; TU 7334 (4), TU 27974 (5), TU 37542 (27) —Gordon Co.; TU 35029 (85)— Walker Co.; TU 27575 ( 1 )— Whitfield Co.; USNM 200704 (4). Middle Coosa River. — Georgia: Chattooga Co.; TU 34978 (2), TU 33367 (3)— Floyd Co.; USNM 31070 (8), USNM 101159 (1), JDS (2), JDS (140). Alabama: Cherokee Co.; JDS (73)— Etowah Co.; USNM 43475 (3)— Cal- houn Co.; TU 40782 (2), UAIC 636 (6), USNM 63136 (4), USNM 200705 (6)— St. Clair Co.; UAIC 648 (6), UAIC 657 (4)— Talladega Co.; UAIC 639 (5), UAIC 880 (1), UAIC 890 (1). Alabama River. — Alabama: Dallas Co.; TU 35357 (39), TU 32640 (28)— Wilcox Co.: TU 40325 (1), TU 2971 (1), TU 3058 (5)— Clark Co.; TU 35344 (1). Cahaba River.— Alabama: Shelby Co.; USNM 43504 (8)— Bibb Co.; UAIC 2035 (1)— Perry Co.; TU 29899 (18), TU 30095 (15), TU 37703 (11), UAIC 962 (49). Tallapoosa Ri\er. — Georgia: Carroll Co.; UAIC 1312 (1). Alabama: Randolph Co.; UAIC 1379 (2)— Elmore Co.; TU 15280 (3), UAIC 1279 (3). Chattahoochee River. — Georgia: Fulton Co.; CU 28249 (2), TU 12167 (1), TU 26125 (1), AU 718 (24)— Heard Co.; AU 512 (3). Acknowledgments I wish to express my appreciation to cura- tors who loaned specimens from their collec- tions: Dr. James E. B5hlke, ANSP; Dr. John S. Ramsey, AU; Dr. H. T. Boschung, UAIC; Dr. Edward C. Raney, CU; Dr. Joseph S. Nelson, lU; Dr. Ernest A. Lachner, USNM; and Dr. W. S. Woolcott, UR. The following persons also extended every courtesy during No. 1 Variation and Relationship of Tuo Studjishes 21 visits to their institutions; Dr. Philip W. Smith, INHS; Dr. Frank B. Cross, KU; Dr. Robert R. Miller, UMMZ; and Dr. Royal D. Suttkus, TU. Drs. Reeve M. Bailey, W. L. Burger, David A. Etnier, Neal Foster, Mr. Robert E. Jenk- ins, Dr. George A. Moore, Mr. William L. Pflieger, Dr. John S. Ramsey, Mr. William Smith- Vaniz, and Mr. James D. Williams all aided the study by supplying color notes, lo- cality data or other useful information. Mr. William L. Pflieger also read the manuscript and offered many helpful comments. Mr. James D. Satterfield kindly supplied con- siderable information from his study of ¥undulns stellifer and allowed me to examine specimens from his personal collection. A number of SIUE students and former stu- dents, particularly Mrs. Norma Brown, Le- land Hill, Dennis Trotter, Steven L. Smith and Gerald L. Smith, helped with various as- pects of the study. Ronald Twillman, Gary D. Waller and Drs. Robert J. and Joyce Goldstein helped make the collections from W Fork Chickamauga Cr. in Georgia. We thank the Goldsteins for their hospitality during this collecting trip. Figure 10 was drawn by Pamela R. Drake. The University Photographic Service assisted in preparing the Figures. Dr. Leonard C. Jones wrote a program and showed me how to operate the GE 265 time sharing com- puter facility at SIUE to treat the data pre- sented in Figures 5 and 6. The study was supported by a Faculty Research Grant from the Graduate School, Southern Illinois Uni- versity. Literature Cited Bailey, Reeve M., (?f a?. 1960. A list Ox com- mon and scientific names of fishes from the United States and Canada. Second Edition. Amer. Fish. Sac. Sp. Piibl. 2:1-102. Bass, J. C, and J. R. Triplett. 1967. An addition to the known Kansas fish fanna. Tram. Kansas Acad. Sci. 7n(3):411. Branson, Braxley A. 1967. Fishes of the Neosho River system in Oklahoma. Am. Mid- land Naturali.'it 78( 1 ): 126-154. CoNANT, Roger. 1960. The queen snake, Na- trix septemvittata, in the Interior Highland of Arkansas and Missouri, with comments upon similar disjunct distributions. Proc. Acad, of Nat. Sci. Philadelphia 112(2):25-40. Cross, Frank B. 1967. Handbook of fishes of Kansas. Mas. Nat. History, Univ. Kansas Misc. Publ. 45:1-357, 20 Figs., 4 color pi. Fowler, Henry W. 1916. Some features of ornamentation in the killitishes or toothed minnows. American Naturalist 50:743-750. CIarman, S. 1895. The eyprinodonts. Mem- oirs Mus. Comp. Zoo/. 19( 1 ):1-179, pis. 1-12. Gerking, Shelby D. 1945. The distribution of the fishes of Indiana. Investigations of Indiana Lakes and Streams 3(1):1-137. Gilbert, Carter R. 1964. The American cyprinid fishes of the subgenus Luxihis (Genus Notropis). Bull. Florida State Mus. 8(2):95-194. Greenwood, P. H. 1964. Environmental ef- fects on the pharyngeal mill of a cichlid fish, Astatoreochromis alluaudi, and their ta.xo- nomic implications. Proc. Linn. Soc. Lond. 176(1 ):1-10. Hall, Gordon E. 1956. Additions to the fish fauna of Oklahoma with a summary of intro- duced species. Southwcitern Nat. 1(1): 16-26. HuBBS, Carl L., and Clark Hubbs. 1953. An improved graphical analysis and comparison of series of samples. Stjst. Zool. 2:49-56, 92. Hubbs, Carl L., and K. F. Lagler. 1947. (2nd printing, 1949). Fishes of the Great Lak?s Region. Bull. Cranhrook Inst. Sci. 26:1-186, pis. 1-26, Figs. 1-251. Mayer, Ernst, G. E. Lixsley and R. L. Usinger. 1953. Methods and Principles of Systematic Zoology. McGraw-Hill Book Co., N. Y., i-ix, 1-328, Figs. 1-45. Miller, Robert R. 1955. An annotated list of the American cyprinodontid fishes of the genus Fundulus, with the description of Fun- clulus per.'iimilis from Yucatan. Occas. Pap. Mus. Zool., Univ. Mich. 568:1-125, 1 pi. Moore, George A. 1968. Fishes, Part II, pp. 21-165, Figs. 1-112. 7»: Vertebrates of the United States (2nd Ed.), by W. F. Blair, A. P. Blair, P. Brodkorb, and F. R. Cagle. McGraw-Hill Book Co., N. Y., 616 pp. Newman, H. H. 1907. Spawning behavior and sexual dimorphism in Fundulus hetcro- clitus and alHed fish. Biol. Bull. 12(5):314- 345, 2 pis. Newman, H. H. 1909. Contact organs in the killifishes of Woods Hole. Biol. Bull. 17(2): 170-180. Ross, Robert D., and J. E. Cahico. 1963. Rec- ords and distribution problems of fishes of the North, Middle, and South Forks of the Holston River, Virginia. Virg. Agr. Exper. Station Tech. Bull. 161:1-24. Smith, Philip W. 1965. A preliminary an- notated list of the lampreys and fishes of Illinois. ///. Nat. Hist. Surv. Biol. Notes. 54: 1-12. Thomerson, Jamie E. 1966. A comparative biosystematic study of Fundulus notatus and Fundulus olivaceus (Pisces: Cyprinodon- tidae). Tulanv Stud. Zool. 13(l):29-47. SWINGLEUS POLYCLITHROIDES GEN. ET SP. N. (MONOGENEA: GYRODACTYLIDAE ) FROM FUNDULUS GRANDIS BAIRD AND GIRARD^ WILMER A. ROGERS Fisheries Laboratory, Agricultural Experiment Station, Auburn University, Auburn, Alabama 36H30 Abstract Swingleus polyelithroides, a new genus and species of gyrodactylid Monogenea, is described from the gulf killifish, Fundulus grandis Baird and Girard. This species is unique in the possession of the following comJMnation of characters, 1) marginal hooks distributed in two groups as in Polyclithrum mugilini Rogers, 1967, 2) a peduncular bar as in Gyrodactylus mega- canthus Wellborn and Rogers, 1967, G. trematoclithrus Rogers, 1967, and G. pro- longis Hargis, 1955, 3) absence of a deep (dorsal) bar, and 4) the presence of lateral accessory bars. The presence of a pedun- cular bar indicates a strong relationship to the species of Gyrodactylus possessing this structure while the marginal hooks being distributed in two groups and the presence of lateral accessory bars shows a relation- ship to Polyclithrum Rogers, 1967. Since the above mentioned features are inter- mediate between the subfamilies Gyrodacty- linae and Polyclithrinae, the subfamily status of Swingletis is unclear at this time. The species described herein was collected as part of a survey of fish parasites being conducted by the Southeastern Cooperative Fish Parasite and Disease Project. Hosts were collected with a seine. The fish were immediately placed in a container of 1:4,000 formalin solution (Putz and Hoffman, 1963). After 1 hour, formalin was added to the con are given followed by the range in paren- theses. Drawings were made with a camera lucida. The keys to the members of Gyro- dactylidae in Yamaguti (1963) were used to determine the status of the present species. Swingleus gen. n. Generic diagnosis: Gyrodactylidae: head bilobed, head organs present. Pharynx py- ramidal, gut bifurcate, not confluent pos- teriorly. Cirrus with spine and several spine- lets. Testis not observed with certainty, appears preovarian. Peduncular bar present. Flaptor with one pair of anchors, ventral bar, pair of lateral winglike bars, and 16 mar- ginal hooks. Dorsal bar absent. Anchor base with cap of dense tissue. Marginal hooks distributed in two groups, each of first three pairs located on anterolateral margin, re- maining five pairs on posterior margin. Parasitic on fish. Type species: Swingleus polyelithroides sp. n. Type host: Fundulus grandis Baird and Girard. Type locality: Mobile Bay at Mobile Causeway, Baldwin County, Alabama. be i Remarks: This genus appears to closely related to Polyclithrum Rogers, 1967 -- ' (Rogers, 1967a) by possessing lateral bars tainer to make a 5% solution. The parasites ^^^ having hooks distributed in two groups, were later recovered for study in the labora- ^^^^^^ shape, ventral bar shape, and the tory from the solution. Specimens were presence of a peduncular bar indicate a strong treated and measured as described by Rogers relationship to certain species of Gyrodacty- and Wellborn (1965). All measurements j^^^ occurring on Fundulus spp., (see Well- are in microns. Averages of measurements ^^^^ ^^^ Rogers, 1967; Rogers, 1967b). Gyrodactylus prolongis Hargis, 1955, was collected at the same time as S. poly- elithroides and anchor and ventral bar shape ' Supported by the Southeastern Cooperative Fish Parasite and Disease Project. ( In part by Sport Fish Restoration funds.) Editorial Committee for this Paper: Dr. John D. Mizelle, Professor of Biological Sciences, Sacramento State College, Sacramento, California Dr. Satyu Yamaguti, Visiting Professor, Laboratory of Parasitology, Department of Biology, Tulane University, New Orleans, Louisiana 22 No. 1 Su'ingleus polycUthroides 23 Figures 1-7. Swingleus polycUthroides gen. et sp. n. Scale of 100 microns applies to Fig. 1; 50 micron scale applies to Fig. 3, 3A; and 20 micron scale applies to all other figmes. 1. Entire worm, ventral view. 2. Anchor. 3. Haptor. 3A. Peduncular bar. 4. Lateral winglike bar. 5. Cir- rus. 6. Hook. 7. \'entral bar and shield. 24 Tulane Studies in Zoology and Botany Vol. 16 of these two species are very similar. A peduncular bar occurs on G. prolongis simi- lar to the peduncular bar of S. polyclithroides. This bar has a series of small ridges and de- pressions appearing as spines. Hargis (1955) reported this bar as an "anteriorly directed skirt whose edge is armed by several cuticu- larized points." The similarities to the above mentioned species of Gyrodactylus (subfam- ily Gyrodactylinae ) and to Polyclithrum ynugilini (subfamily Polyclithrinae) makes the subfamily status of S. polyclithroides un- clear at this time. This genus is named in honor of Dr. H. S. Swingle of Auburn Univer- sity. Sivingleus polyclithroides sp. n. (Figures 1-7) Host and locality: Fundulus grandis Baird and Girard, Gulf killifish. Mobile Bay at Mobile Causeway, Baldwin Co., Alabama. Location on host: Fins and body. Specimens studied and measured: Ten. Type specimens: Holotype and 2 para- types, USNM Helm. Coll. Nos. 70445 and lOAAG; paratypes in author's collection. Description: Length 479 (390 to 620), width 94 (80 to 130). Cephalic area with two prominent lobes, each containing several spines. Cephalic glands prominent, lateral to posterior edge of pharynx; head organs well developed. Pharynx bilobed in side view, posterodorsal lobe larger, 35 (30 to 39) in diameter, anteroventral lobe 28 (25 to 39) in diameter. Gut bifurcate with crura end- ing blindly at level of gonads. Cirrus ( Fig. 5 ) located ventrally to left of and posterior to pharynx, with large spine and 2 to 5 spinelets, diameter 15 (13 to 19). Testis preovarian? Ovary with many diffuse eggs in various stages of development. Peduncular bar present, with many small oblong to elon- gate depressions (Fig. 3A), greatest length of bar 66 (63 to 69); width 19 (18 to 20). Haptor (Fig. 3) circular to oblong, with one pair of anchors supported by a ventral bar and lateral "winglike" bars, and 16 marginal hooks. Haptor length 109 (105 to 135), width 96 (70 to 108). Anchors (Fig. 2) with prominent knobs apparently represent- ing vestigial dorsal bar, points with arc mem- branes (Mizelle and Kritsky, 1967) and small membranes arising from shaft near point similar to "wings" of dactylogyrids, base of anchor shaft with dense cap of tissue; anchor length 96 (92 to 100), width 10 (9 to 11), point length 23 (21 to 25 ) . Ventral bar ( Fig. 7 ) articulated in prominent folds on anchor shaft, with shield terminating at posterior edge of haptor, anterolateral projec- tions of ventral bar with thickened area, cen- tral portion of bar with coarse striations. Bar 37 ( 33 to 45 ) by 11 (9 to 12), antero- lateral projection length 12 (10 to 13). Shield of ventral bar with coarse striations on margins and 2 forked ridges postero- medially, shield length 46 (42 to 48), width 42 (40 to 45). Lateral bars (Fig. 4) wing- like, with coarse striations distally, base ar- ticulated to anchor shaft, length along bar axis 37 (34 to 39). Marginal hooks 16 in number, located dorsally on haptor, first 3 pairs in 2 groups at anterolateral edge of haptor, remaining 5 pairs on posterior edge of haptor (Figs. 1, 3). Shaft of each hook ( Fig. 6 ) with slight enlargement proximally, shaft length 32 (30 to 33), hook length 7 to 8, lamella length 12 (11 to 13). Remarks: This is the only species reported in the genus. It was found on the host in association with Gyrodactylus prolongis Har- gis, 1955. The name polyclithroides refers to the similarity to the genus Polyclithrum. Acknowledgments Thanks are extended to Mr. John R. Kel- ley, who initially collected this species, and to Messrs. Shih-Ming Chien, S. K. Johnson, and M. V. Rawson for subsequent collections. Dr. John S. Ramsey confirmed host identifi- cations. Literature Cited Hargis, W. J. 1955. Monogenetic trematodes of Gulf of Me.xico fi.shes. Part 1, The Super- family Gvrodactyloidea. Biol Bull 108:125- 137. PuTz, R. E., and G. L. Hoffman. 1963. Two ne-w Gyiodacttihis (Trematoda: Monogenea) from c>prinid fi.shes with a synopsis of those found on North American fishes. J. Parasit. 49:559-566. Mizelle, J. D., and D. C. Kritsky. 1967. Studies on monogenetic trematodes. XXX. Five new species of Gtirodactrihis from the Pacific Tomcod, Microgadti.s pruxiinus (Gi- rard). /. Parasit. 53:263-269. No. 1 Swingleus polyclithroides 25 Rogers, W. A. 1967a. Polyclithrum mugilini gen. et sp. n. (Gyrodactylidae: Polyclithrinae subfani. n. ) from Mtigil cephalus L. /. Para- sit. 53:274-276. . 1967b. Six new species of Gyrodacttj- his (Monogenea) from the Southeastern U.S. /. Parasit. 53:747-751. and T. L. Wellborn, Jr. 1965. Stud- ies on Gtjrodactyhis (Trematoda: Monogenea' with descriptions of five new species from the Southeastern U.S. /. Parasit. 51:977-982. Wellborn, T. L., Jr., and W. A. Rogers. 1967. Five new species of Gyrodactyhis (Trematoda: Monogenea) from the Southeastern U.S. /. Parasit. 53:10-14. Yamaguti, S. 1963. Systema Hehninthum Vol. IV. Monogenea and Aspidocotylea. Inter- science Publishers. New York and London. December 16, 1969 A NEW TREMATODE, LEPIDODIDYMOCYSTIS IRW^INI N. G., N. SP. (DIDYMOZOIDAE) FROM A MARINE FISH, MENTICIRRHUS NASUS SATYU YAMAGUTI and SHUNYA KAMEGAI Lahoratonj of Parasitology, Department of Biology, Titlane University, New Orleans, Louisiana 70118 Abstract A new didymozoid belonging to Didy- mozoinae Ishii, 1935, is described. It is named after its collector, Mr. Roy Irwin, a graduate student at Tulane University. It was found underneath the scales below the lateral line of a marine teleost, Menticirrhus nasits (Gunther, 1868), from N. Mazatlan. The generic name refers to this special hab- itat. The most outstanding features of this trematode include: (1) flat semicircular hindbody with a distinct longitudinal fur- row on concave side, (2) acetabuhun pres- ent, (3) common genital pore opening at truncate apex of prominent genital papilla, (4) long esophagus, ( 5 ) single imdivided ovary and single undivided \itelline gland, (6) cylindrical sinuous egg reservoir, (7) metraterm ciliated throughout its length, and ( 8 ) eggs not embryonated when laid. Introduction Twenty specimens on which the present report is based were taken by Mr. Roy Ir- win, a graduate student at Tulane University, together with scales from below the lateral line of Menticirrhus nasus collected in N. Mazatlan, Mexico. They had been previously fixed in for- malin in situ; they were isolated from the cysts attached underneath the scales, refixed in 10% formalin, and stained with Dela- field's hematoxylin. We gratefully acknowl- edge Mr. Irwin's furnishing the material for our examination. The worms, enclosed in pairs in transparent circular cysts, were so strongly flattened underneath the scales that the whole mounts clearly revealed the inter- nal anatomy, making sectioning unnecessary. This parasite represents an undescribed didy- mozoid for which Lepiclocliflyinocystis irwini n. g., n. sp. is proposed. Description Based on 20 whole mounts. Cysts flattened circular, about 3-5 mm in diameter. Fore- body flattened claviform, conspicuously papil- lated at head end, finely annulated for re- maining part, 0.53-0.82 mm long with maxi- mum width of 0.13-0.32 mm posterior to its middle, whence it tapers gradually forward to a more or less sharp point, attached to anterior end of hindbody on its concave side. Hindbody flattened, semicircular, truncate anteriorly and rather pointed posteriorly, 3.6-5.1 X 2.44-3.58 mm; forebody embedded in a distinct longitudinal furrow on concave side of hindbody. Oral sucker terminal, ob- long, 66-82 X 43-52 /x, weakly muscular, di- rectly followed by spherical muscular pharynx 42-49 jx in diameter. Immediately behind pharynx clustered large claviform pharyngeal gland cells, with attenuated ends converging toward posterior end of pharynx. Esophagus narrow, 0.46-0.87 mm long, provided with a coat of small glandular cells throughout its length, bifurcating immediately as it enters hindbody; ceca lined with epithelia, strongly winding throughout length of hindbody and terminating at its posterior end. A cup- shaped, weakly muscular acetabulum 59-91 /x in diameter lies on concave side of hind- body near base of forebody. Testes paired, long, tubular, winding, usually close to each other, rarely divergently in anterior part of hindbody near concave side; in type they reach just beyond equatorial level, with their Editorial Committee for this Paper: Dr. J. Teague Self, Regents Professor of Zoology, University of Oklahoma, Nor- man, Oklahoma Dr. Allen McIntosh, 4606 Clemson Road, College Park, Maryland 26 No. 1 Lepidodidymocystis irwini 21 ends somewhat swollen. Vas deferens nar- that the genus in question belongs in the row throughout its length, without forming Didymozoinae. It is named in reference to a definite seminal vesicle, distinctly ciliated the special habitat (beneath the scales), and just before uniting with metraterm at base defined as follows: of genital papilla. Genital papilla projecting GENERIC DIAGNOSIS. Didymozoidae, Didy- prominently ventral to oral sucker, 25-40 /x mozoinae. Complete hermaphrodites, cn- in diameter, flattened at apex where wide cysted in pairs. Forebody small, flattened common genital pore opens. Ovary a single, claviform, attenuated anteriorly, attached to long, narrow tubule, 23-53 /<. wide, irregu- hindbody near its anterior end, papillated at larly winding in axial region of hindbody, head end, and finely annulated elsewhere, reaching to near its posterior end. Vitelline Hindbody smooth, approximately scmicircu- gland also a single, irregularly winding, nar- lar, truncate at anterior end, conical at pos- row tubule, 30-67 \x. wide, extending on con- terior end, with distinct longitudinal furrow vex side of hindbody to its extreme posterior on concave side, where a cup-shaped, weakly end, its anterior portion turns back on itself muscular acetabulum is present near base of at truncate anterior end of hindbody and forebody. Oral sucker terminal, weakly mus- after describing several turns joins proximal cular; pharynx spherical, muscular, with well end of ovary. This end is often swollen like developed postpharyngeal gland cells behind, proximal end of vitelline gland; seminal Esophagus long, narrow, surrounded by small receptacle 74-178 X 45-97 /a, situated at this glandular cells throughout its length, bifur- genital junction. Uterine duct provided with eating as it enters hindbody. Ceca strongly a thick coat of gland cells, runs sinuously winding, terminating at posterior extremity forward to recurrent portion of vitelline of hindbody. Testes two, tubular, winding in gland, where it passes to uterus proper, the anterior half of hindbody close to concave latter winding backward to posterior end of side; vas deferens narrow, not forming defi- the hindbody and then forward to anterior nite seminal vesicle, ciliated just before unit- end of hindbody and once more backward, ing with metraterm. Common genital pore to lead into t^g reservoir near the posterior opening at apex of prominent truncate geni- extremity, thus forming four loops altogether tal papilla ventral to oral sucker. Ovary sin- and occupying greater part of hindbody; (igg gle, tubular, long, undivided, winding in reservoir cylindrical, sinuous, close to con- axial region of hindbody and reaching to cave side of hindbody; metraterm well dif- near its posterior end. Vitelline gland tubu- ferentiated in forebody, distinctly ciliated lar, long, undivided, winding from extreme inside throughout its length. Egg oval, thick- posterior end of hindbody to its anterior end, shelled, operculate, 12-15 X 9-11 /a, ova con- where it turns backward to join anterior end tained in metraterm not yet embryonated, of ovary, so that genital junction lies near almost all in 2-cell stage. Excretory system truncate anterior end of hindbody. Seminal not made out. receptacle present. Uterus occupying all P^ available space of hindbody, forming four longitudinal loops before leading into con- In general internal anatomy and habitat spicuous egg reservoir which extends longi- this genus bears a marked resemblance to t^jjnally along concave side of hindbody; Dermatodtdyvwcysus Yamaguti (especially ^^,^^^^,^ ^ell differentiated, ciliated inside D. viviparoides Yamaguti) (in press) but , , • i u t- i . ..u.vi j-rr r • ■ 1 • c ^. throughout Its length. Eggs operculate, thick- dirters from it in the possession of : ( 1 ) a , ,, , , , . ^ „ longitudinal furrow and an acetabulum on shelled, not embryonated tn utero. Excretory the concave side of the hindbody, (2) a system not made out. Parasitic underneath prominent genital papilla ventral to the oral scales, especially below lateral line, of marme sucker, ( 3 ) a uterus regularly forming four teleosts. longitudinal loops before leading into a Type species: Lepidodidymocystis irwini prominent egg reservoir, and (4) unem- n. g., n. sp. in Menticirrhus nasus\ N. bryonated eggs, an important character since Mazatlan. Forebody 0.53-0.82 X 0.13-0.32 most other didymozoid eggs are generally mm, hindbody 3.6-5.1 X 2.44-3.58 mm; eggs embryonated in utero. There is no doubt 12-15 X 9-11 /i. 28 Tulane Studies in Zoology and Botany Vol. 16 No. 1 Lepidodidymocystis irivini 29 Literature Cited Yamaguti, S. Systema Helminthum, Vol. 1, re- IsHii, N. 1935. Studies on the family Dicl>- \isecl and enlarged. (In press.) niozoidae ( Monticelli, 1888). Jap. J. Zoo/. . Digenetic Treinatodes of Hawaiian 6(2):279-335. fishes. (In press.) December 16, 1969 Figures 1 to 4. Lepidodidymocystis incini n. g., n. sp. Fig. 1. Holotype, lateral view; Fig. 2. Paratype 16, lateral view of entire forebody and part of hindbody; Fig. 3. Paratype 7, general \iew of forebody and acetabulum; Fig. 4. Paratype 7, \entral \ iew of anterior extremit\- of fore- body. Abbre\'iations used in figures: A acetabulum; C cecum; E esophagus; ER egg reservoir; F lon- gitudinal furrow on concave side of hindbody; FB forel)<)d>-; C.P genital pore; HB hindbody; M metraterm; N nerve; O ovary; OS oral sucker; P pharynx; T testis; U uterus; VD vas deferens; NT vitellarium. > i Volume 16, Number 2 February 25, 1970 MUS. COMP. ZOCL. LIBRARY \m 1 0 1970 HARVARD U^aVERSlTYl THE COACHWHIP SNAKE, MASTICOPHIS FLAGELLUM (SHAW): TAXONOMY AND DISTRIBUTION LARRY DAVID WILSON Department of Biology, University of Southwestern Louisiana, Lafayette, Louisiana 70501 p. 31 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. 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Haddenhorst Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 16, Number 2 February 25, 1970 THE COACH WHIP SNAKE, MASTICOPHIS FLAGELLUM (SHAW): TAXONOMY AND DISTRIBUTION^ LARRY DAVID WILSON Department of Biology, Uiiivcrsitij of Sotithwestern Louisiana, Lafayette, Louisiana 70501 CONTENTS Abstract 31 Introduction 32 Historical Summary 33 Validity of the Genus Mastkophis 33 The flagelhwi Group of Mastkophis 34 Composition of the Species Mastkophis flagellnni 35 Subspecies Accounts 36 Masticophis flagellum jlagelhmi ( Shaw ) 36 Masticophis flagellum testaceus ( Say ) 42 Masticophis flagellum lineatidus Smith 47 Masticophis flagelh/m picens (Cope) 51 Masticophis jlagellmn ruddocki Brattstrom and Warren 55 Masticophis flagellinn cingidiim Lowe and Wood 58 Masticophis flagelhmi fidiginosKS (Cope) 63 Key to the Subspecies of Adult Masticophis flagellum 73 Significance and Evolution of Color Pattern _. 74 Variation in Scuttelation and Proportions 77 Distribution and Habitat 89 Summary and Conclusions 93 Acknowledgments 94 Literature Cited 95 Abstract the species flagcUtmi to the otlier species within tlie genus Maslieophis are hriefly The taxonomy and distribution or Mas- considered. ticophis flagellum (Shaw) are discussed. 5i^ subspecies recognized previously by The use of the generic name Masticophis ^t^gr authors are retained, and fuUgim)sus instead of Coluber and the relationship of Cope is resurrected from synonymy for another subspecies. The concept of the 1 This work was completed in partial fulfill- subspecies cingulum is modified. A dis- ment for the degree of Doctor of Philosophy at cussion of color pattern \ariation is in- Louisiana State University, Baton Rouge, Lou- eluded under each subspecies heading, isiana. The tjpes of variation illustrated !)>• the Editorial Committee for This Paper: Mr. Roger Conant, Director and Curator of Reptiles, Philadelphia Zoological Gar- den, Philadelphia, Pennsylvania 19104 Dr. Albert Schwartz, Department of Biology, Miami-Dade Junior College, Miami, Florida 31 32 Tnhme Studies in Zoology and Botany Vol. 16 color pattern are discussed, and so are infraspecific relationships. The \ariation exhibited by the nonpat- tern characteristics is summarized and analyzed for se.xual, ontogenetic, geo- graphic, and indi\'idual significance. Available information on distribution and habitat ijreference is reviewed. Introduction The coachwhip snake, Masticophis flagel- lum, is a wide-ranging North American species that exhibits considerable pattern variation. It has not been reviewed in its entirety since Ortenburger's (1928) revi- sion of the genera Masticophis and Coluber. Ortenburger had approximately 400 speci- mens of M. jlagellu?7i available to him; in contrast I have examined more than five times as many. I recognize seven sub- species, three of which have been described since 1928. I have studied variation in most of the characters normally utilized in taxo- nomic work on snakes at the specific level, with emphasis on color pattern. These char- acters have been analysed for possible geo- graphic, sexual, ontogenetic, and individual differences. During the course of this study I have examined 2195 specimens of Masticophis flagellum from the following institutional and private collections: AC Alabama College AMNH American Museum of Natural History ANSP Academy of Natural Sciences of ASDM Philadelphia Arizona-Sonora Desert Museum AU Auburn University BCB Biyce C. Brown Collection, Strecker Museum, Baylor University BYU Brigham Young University CAS California Academy of Sciences CHM Charleston Museum CM Carnegie Museum CU Cornell University DEH Donald E. Hahn, private collection EAL Ernest A. Liner, private collection FSU Florida State University HC Howard College INHS Illinois Natural History Survey JFC Joseph F. Copp, private collection JPC Janalee P. Caldwell, private collec- tion JRD James R. Dixon, private collection KLW Kenneth L. Williams, private collec- tion KU University of Kansas Museum of Natural History LACM Los Angeles County Museum LPI Louisiana Polytechnic Institute LSUMZ Louisiana State University Museum of Zoology MCZ Museum of Comparative Zoology MGFCM Mississippi Game and Fish Commis- sion Museum MMH M. Max Hensley, private collection MISU Mississippi State University MSU Michigan State University NCSM North Carolina State Museum NLSC Northeastern Louisiana State College NMSU New Mexico State University NSU Nevada Southern LIniversity NTSU North Texas State University OSU Oklahoma State University PM Philip Medica, private collection RAT RoJjert A. Thomas, private collection SDSNH San Diego Society of Natural His- tory SM Strecker Museum TCWC Texas Cooperative Wildlife Collec- tion TNHC University of Texas Natural History Collection TU Tulane University UAHC University of Alabama Herpetologi- cal Collection UAZ University of Arizona UCM University of Colorado Museum LIE University of Florida Collections UG University of Georgia UIMNH University of Illinois Museum of Natural History UM University of Miami UMMZ University of Michigan Museum of Zoology UNM University of New Mexico USL University of Southwestern Louisi- ana USM University of Southern Mississippi USNM United States National Museum UU University of Utah Scale counts were made in accordance with customary methods. Ventrals were counted using the method of Dowling (1951b), and so were scale reductions (Dowling, 1951a). Because of extreme vari- ability in the temporal series, the number of scales on each side of the head was totaled as a single number. Snout-vent length was measured from the tip of the snout to the posterior edge of the anal plate. Tail length was measured from the posterior edge of the anal plate to the tip of the tail. I Unless otherwise noted, the color pattern descriptions are based on preserved material, although I have made use of whatever living material was available. Brackets are used in the text to indicate where a qualifying statement of mine has been made within a quotation. I have used brief synonymies for the sub- No. 2 Coachichip Snake, Masticophis flagellmn 33 species discussed by Ortenburger (1928) and complete synonymies for those sub- species described since Ortenburger's work. Historical Summary Coluber flagellum was described by Shaw ( 1802 ) from a plate in Catesby's "The Nat- ural History of Carolina, Florida and the Bahama Islands." The provenance of the species was given as North America, and Shaw stated that it is "not uncommon in Carolina and Virginia." These states were regarded as the type locality by Ortenburger (1928). In 1823 Say described CoJiiber testaceus from the western portion of the range. Hallowell (1852) described Psani- fuophis flai'igularis from Cross Timbers, Oklahoma. The specific name flavigularis was used by Ortenburger for the western coachwhip, instead of testaceus, but most investigators have considered fhwig/daris to be a synonym of testaceus. Several subspecies have been described. Stejneger (1893) added Bascanion flagellum jrenatmn, from Mountain Spring, San Diego County, Cali- fornia. Masticophis flagellum lineatulus was described by Smith in 1941 from 11 miles north of San Buena Ventura, Chihuahua, Mexico. Masticophis flagellum ruddocki was described by Brattstrom and Warren in 1953 from Kern County, California. Lowe and Woodin added a sixth form, Masticophis flagellu7)i ci}!gulu7u, in 1954, based on speci- mens from Moctezuma, Sonora, Mexico. Ortenburger (1928) was the first investi- gator to review the genus, and, in his excel- lent paper, he recognized three subspecies of flagellum {flagellum, flavigularis, and frena- ti/m ) . He considered piceus, a name for- merly applied to black individuals of frena- ttivi, as a distinct species. But much more material has since accrued and three addi- tional subspecies have been described. It is now possible to outline the range of each race with considerable accuracy, and to de- lineate the areas of intergradation between subspecies. Also, the considerably greater amount of specimens now available makes it possible to analyze the variation within the species with greater accuracy. Validity of the Genus Masticophis Considerable controversy exists in the literature as to whether or not the snakes grouped under Masticophis should be con- sidered as generically distinct from the snakes of the genus Coluber. In his mono- graph, Ortenburger (1928) discussed what he considered to be the distinguishing char- acteristics between Masticophis and Coluber. These included differences in the number of maxillary teeth, the type of scale reduction formula, the number of ventrals, subcaudals, and supralabials ( including the number of supralabials entering the orbit, usually a function of the number of supralabials), certain characteristics of hemipenial struc- ture, and the head width-head length ratio. The severest critics of the use of the generic name Masticophis were Bogert and Ohver (1945). They cited Stuart's (1934) difficulty in deciding whether his new species ortenburgeri belonged to the genus Masticophis or Coluber. Stuart stated that the type specimen agreed with members of the genus Masticophis in having a similar scale reduction pattern, but, on the contrary, it agreed with members of the genus Colu- ber in its hemipenial structure. Coluber or- tenburgeri has recently been shown to be the southernmost representative of and synonymous with Coluber constrictor oaxaca i=C. c. stejnegerianus auct. ) (See Eth- eridge, 1952; Stuart, 1963; Wilson, 1966). In addition, during the preparation of my paper on Mexican Coluber ( op. cit. ) , I examined the holotype of Coluber orten- burgeri and found it to be somewhat aber- rant. The complete scale reduction in this specimen is: 3+4(5) 3 + 4(70) 6 + 7(129) 19 17 15 3 + 4(5) 14 3 + 4(70) 6 + 7(139) 7 (129) 15 14 This specimen has neither a typical Coluber nor a typical /Masticophis scale row pattern, but it more closelv resembles Coluber con- strictor. Dunn (1933) believed that the snecimen of Coluber oaxaca ( — C. c. oaxaca) from Colima, Mexico that he discussed demonstrated a similarity to Masticophis in scale row reduction and number of supra- labials. As I have stated previously (Wil- son, 1966, p. 45), I believe this scale re- duction pattern to be aberrant. The num- ber of supralabials (eight) is, indeed, the 34 Tulane Studies in Zoology and Botany Vol. 16 same as that seen in Masticophis, but eight supralabials is also characteristic of the west- ern forms of Coluber constrictor (i.e., mor- 7non and oaxaca); seven is characteristic of the more eastern forms. The mean number of supralabials increases gradually from east to west. Populations of Coluber constrictor in the eastern portions of Louisiana, for example, have a lower mean number of supralabials than those from the western portions of the same state (Wilson, unpub- lished manuscript). Use of characters ex- hibited in limited portions of the range of Coluber constrictor would not appear to be a valid criterion for demonstrating that Masticophis and Coluber are congeneric, nor are aberrant characters. Inger and Clark's (1943) attempt to sub- divide the cumbersome genus Coluber ( sensu lato) has also met with criticism. Bogert and Oliver (1945) stated that these authors "attempt to partition the genus Coluber (in its broadest sense) on the sole basis of scale reduction patterns." This is not strictly the case because Inger and Clark used the num- ber of supralabials entering the orbit and the presence or absence of enlarged basal spines on the hemipenis as supportive char- acters. Although I would agree that the sub- division of or the erection of a generic taxon, if based on a single character, is ill advised, I also believe that Inger and Clark's system contributes to a better understanding of the relationships of this group of snakes. As I have stated (Wilson, 1967, p. 269), the genus Coluber ( sensu lato ) is an unwieldy assemblage of snakes, many of which are only distantly related to and are even gen- erically distinct from Coluber constrictor, the type species of the genus. The range of characters exhibited collectively by the spe- cies now grouped in the genus Coluber is very wide and indicative of an unnatural grouping. The Coluber complex admittedly needs further study, and the relationships of its many components may not become clear until investigations are made on skele- tal anatomy, hemipenial structures, blood proteins, and behavior on a world-wide basis. To retain the species of the genus Masticophis in the already unwieldy Coluber would require inclusion also of the other New World racer genera, i.e., Dryadophis, Drymobius, Leptodry?nus, and Dendrophi- dion, which are certainly less distinct from Masticophis and Coluber constrictor than are the Old World Coluber dorri and Colu- ber dipsas. For these reasons I continue to use the name Masticophis for the whip- snakes in general and for the species flagel- lum in particular. The flagellum Group of Masticophis In his review of Masticophis Ortenburger ( 1928) divided the genus into two sections, the taeniatus group and the flagellum group. His taeniatus group included M. aurigulus, M. barbouri, M. lateralis, M. ruthveni ( = M. taeniatus ruthveni), M. schotti ( = M. taeniatus schotti), M. semilineatus { — M. bilineatus) , M. taeniatus girardi, and M. taeniatus taeniatus. This group is not perti- nent to the present study and will not be discussed further. Ortenburger's flagellum group included M. anthonyi, M. flagellum flagellum, M. f. flavigularis {— M. f. testaceus ) , M. f. frenatum {— M. f. piceus) , M. linea- tHs {— M. striolatus), M. mentovarius, and M. piceus (= M. f. piceus). This arrangement has seldom been challenged, and I see no reason for changing it, except to shift some names because of priorities. Smith (1941), who discussed the Mexican forms of the genus, presented a phylogeny that did not differ significantly from that of Ortenburger insofar as the division of the constituent species into two groups is con- cerned. Smith, however, suggested that mentovarius is similar to the basic stock from which the remainder of the species developed. Ortenburger, on the other hand, postulated that the two groups arose from a " . . . hypothetical progenitor of the genus Masticophis y It is my opinion that a proto- ?nentovarius was not the basic form and that, furthermore. Smith's phylogeny represents the relationships of the species of the genus Masticophis at the present time level. Without adequate paleontological data it is futile to speculate on the age of certain species, but it is more logical to assume a progenitor of the genus that diverged to form two groups: (1) a banded or unpat- terned form (the flagellum group), and (2) a striped form (the taeniatus group). Masticophis mentovarius appears to be a derivative of striolatus that migrated south- ward to invade Central and South America. It is likely that the peculiar fusion of the No. 2 Coachwhip Snake, Masticophis jlagellum 35 F'igiire 1. Masticophis jlagellum jlagellum (CHAS 6722) from Vaiden, Carroll County, Mis- sissippi. fourth and fifth supralabials is a derived character, inasmuch as all the other members of the genus have two supralabials entering the eye, as do the members of the genus Coluber. A similarity in pattern ties 7nentovarws to striolatns. Most likely, another derivative of striolatus is M. authonyi, found only on Isla Clarion in the Revillagigedo Island group ( Ortenburger, 1928; Brattstrom, 1955). These three species form a compact group that is distinct from all the subspecies of Masticophis jlagellum, except for lineatu- liis, in having a lineate dorsal pattern com- posed of a stripe (or a dot in the case of anthonyi) on each scale. The pattern of lineatulus, however, appears to have been derived separately from that of the striola- tus subgroup, because the juveniles are typi- cally banded across the neck, as is the case in the young of the other subspecies of \lagellum. Juveniles of lineatulus are identi- cal with juveniles of M. f. testaceus in this respect. Composition of the Species Masticophis flagellu??i Smith and Taylor (1945) listed six sub- species of Masticophis jlagellum ( five Mexi- can subspecies and one extralimital form, M. j. jlagellum ) . Since that time two ad- ditional subspecies {cingulum and ruddocki) have been described, and two subspecies have been removed. Bogert and Oliver (1945) demonstrated sympatry between Masticophis striolatus ( = Coluber striolatus striolatus of Bogert and Oliver and Mastico- phis jlagellum lineatus of Smith and Taylor) and Masticophis jlagellum cingulum ( = Co- luber jlagellum piceus of Bogert and Oliver and Masticophis jlagellum piceus of Smith and Taylor ) in southern Sonora. They also stated that "the snake described by Smith (1943, p. 448) as Masticophis jlagellum variolosus, if it proves to be valid, should be placed as a subspecies of Coluber strio- latus." Zweifel ( I960), in his study of the herpetofauna of the Tres Marias Islands, dis- cussed the validity of Smith's variolosus and concluded that "... differentiation in num- ber of ventral scales alone is not an adequate basis for the recognition of an endemic in- sular subspecies. Whether variolosus should be recognized or not depends on the signifi- cance attached to slight but possibly con- sistent difference in lip pattern, in the apparent absence of more consistent differ- ences. To recognize the subspecies would place undue emphasis on slight variation in a species not otherwise subject to variations sufficiently consistent for it to be considered polytypic, unless it proves to be conspecific with M. mentovarius. Hence, variolosus should be relegated to the synonymy of lineatus'.' Webb (I960) presented evidence confirming the specific status of M. strio- latus. Further confirmation has been pre- sented by Fugler and Dixon (1961) and Smith and Van Geldcr ( 1955). Thus, Masticophis jlagellum presently contains six subspecies {cingulum, jlagellum, lineatulus, piceus, ruddocki, and testaceus). Masticophis striolatus is now recognized as a distinct monotypic species. There remains, 36 Tiilane Studies in Zoology and Botany Vol. 16 however, the possibiUty that striolatus may be conspecific with M. mentovarius (Webb, I960; Zweifel, I960). Subspecies Accounts Masticophis flagellum flagellum (Shaw) Eastern Coachwhip Coluhcr flagellum Shaw, 1802: 475 Coluber testaceus: Harlan, 1826: 348 Psammophis flagelliformis: Holbrook, 1842: 11 Masticophis flagelliformis: Baird and Girard, 1853: 98 Masticophis flavigularis: Baird and Girard, 1853: 99 (part) Herpetodryas flagelliformis: Dumeril and Bi- bron, 1854: 210 Herpetodryas flavigularis: Giinther, 1858: 118 (part) Masticophis flagelliformis testaceus: Jan, 1863: 65 (part) Bascanium flagelliforme flagelliforme: Cope, 1875: 40 Bascanium flagelliforme testaceum: Yarrow, 1875: 542 (part) Bascanium flagelliforme: Cope, 1877: 64 Bascanion flagelliforme: Smith, 1882: 636 Coluber flagelliformis testaceus: Garman, 1883: 43 (part) Bascanion flagelliformis testaceus: Dumeril and Bocwirt, 1886: 705 Zamenis flagellifortnis: Boulenger, 1893: 389 ( part ) Zamenis flagellum flagellum: Cope, 1900: 789 ( part ) Zamenis flagelliforme: Ditmars, 1912: 218 Coluber flagellum flagellum: Stejneger and Barbour, 1917: 79 (part) Masticophis flagellum flagellum: Ortenburger, 1923: 2 Holotype. — None designated. The original description by Shaw (1802) consisted pri- marily of information supplied by Catesby (1731-43). Shaw stated "it is a native of North America, and not uncommon in Caro- lina and Virginia." Schmidt (1953) re- stricted the type locality to Charleston, South Carolina. Diagnosis. — A subspecies of Masticophis flagellum characterized by a dark to very dark brown anterior dorsal coloration and a tan to dark brown posterior dorsal colora- tion. The venter is dark brown anteriorly, grading to cream posteriorly ( Fig. 1 ) . Range. — Southeastern United States from North Carolina south to and including all of Florida, west to eastern Texas, north to Missouri and southern Illinois, including the eastern half of Oklahoma and Kansas ( Fig. 2). Description. — The only information given by Shaw (1802) was that this snake "is a very long and slender species, measuring from four to six feet or more in length: its color is a uniform dusky brown, palest be- neath." The following color pattern description is from a live female specimen (LSTJMZ 9679) collected beside the Red River at the south city limits of Alexandria, Rapides Parish, Louisiana and measuring 1029 mm in head-body length. The anterior portion of the dorsum is very dark velvety brown. This color grades into pinkish brown on the posterior portion of the body and tail. Along the middle of the body, at intervals of about 100 mm, are a series of light bands about 40 to 50 mm long. They are orange brown dorsally and brownish gray laterally. There are about three or four of these bands, the last of which gives rise to a middorsal stripe-like area that is somewhat lighter than the lateral area. The venter is grayish brown on the neck with light orange flecks grading to cream posteriorly with an over- tone of orange flecks. This orange colora- tion is darker on the lateral edges of the ventrals. The underside of the tail is pinkish orange. The head is dark velvety brown above. There is an orange spot on the right parietal. The lateral part of the head is grayish brown anteriorly, becoming brown on the temporals. There is a light spot on the upper preocular. The chin is grayish brown with white blotches on all scales. There are a few orange flecks on the gulars. The iris is dark brown except for an orange ring around the pupil. Discussion. — Meristic and mensural data for this subspecies, based on 241 males and 207 females, may be summarized as fol- lows: supralabials 7 to 9, infralabials 8 to 13, preoculars 2 or 3, postoculars 1 to 3, loreal 0 to 3, ventrals in males 188 to 212, ventrals in females 186 to 207, subcaudals in males 100 to 122, subcaudals in females 91 to 123, dorsal scale rows 19-17-11 to 19-17-15. Body length in males of all ages ranges from 291 to 1600 mm, in females of all ages 331 to 1520 mm. Total length in males of all ages ranges from 373 to 2074 mm, in females of all ages 436 to 1983 mm. Tail length /total length ratios in males ranges from 0.220 to 0.275, in females 0.217 to 0.300. There is a considerable amount of pattern No. 2 Coachwhip Snake, Masticopbis flagellum 37 «•' *••••. a • •• „ o^ .• •••• '• . -^^'' 0) CO (L> a o C a cS X d) CO c 0) O 0) CO O CS o o c CO o u CO lU u CS ♦J <£ tso o f*^ *+-% 60 •** .« a . o V5 t^ •*-* CO a a :§ o <-lH o tu ft c >> _o 4-' x •^ u -C Cw 'C v: *J CO ^ Q ci \r. T3 (U ^ ^ C 3 O M 0) u E i> 38 Tulane Studies in Zoology and Botany Vol. 16 variation in the nominate subspecies. This subspecies exhibits an ontogenetic color change. Juveniles of M. f. flagellum have a pattern similar to the young and adults of M. f. testaceus. The dorsum is some shade of tan with narrow brown to dark brown bands on the neck and forebody, which diminish gradually in intensity posteriorly. The head is light brown or tan, with white outlining the dorsal scales, and with a dark brown area on the posterior portion of the internasals and prefrontals, on most of the supraoculars and frontal, and on the anterior, lateral, and posterior portions of the parie- tals. The side of the head is light brown to tan with cream spots on most scales. The chin is cream, with light brown pigment between the junction of the infralabials and the chin shields. The venter is uniforrp cream, except for a double row of brown spots on the neck. As the snake increases in size the dorsal pattern on the anterior part of the body be- comes uniformly dark as the result of a darkening of the existing brown or tan coloration on each scale and the invasion of the pale lateral areas of each scale by the same color. The dorsal surface of the head becomes uniformly dark by loss of the white scale borders and by a darkening of the light areas on the head. The side of the head becomes darker in an anterior to pos- terior direction. The chin becomes uniformly dark by the deepening of the existing brown color and a progressive invasion of that color into the light cream areas. The spots situated on the anterior portion of the ven- ter begin to darken and enlarge. As the snake grows older and larger, these spots fuse together and still later they fuse with the lateral extension of the dorsal ground color to form the uniformly dark, anterior ventral coloration of the adult. This description of the ontogenetic color pattern changes in flagellum is generalized; some individuals of this subspecies may reach a certain level in the development of the color pattern at a smaller or larger size than others. For example, in LSUMZ 16772 from Caddo Parish, Louisiana (625 mm in snout-vent length), and LSUMZ 16773 from Sabine Parish, Louisiana (621 mm), the former is much paler dorsally than the latter. The juvenile markings are still evident on the head of LSUMZ 16772 and the spotting on the venter is more clearly de- fined. These two snakes were collected only a few miles apart. A juvenile flagellum (OSU 483) with a curious pattern is available from Lake Carl Blackwell in Payne County, Oklahoma. This specimen, which measures 540 mm in total length, entirely lacks the typical neck bands. Instead, each of the dorsal scales, especially on the anterior portion of the body, has a concentration of darker pigment toward the center of the scale, giving the impression of a broad, diffuse stripe on each dorsal scale row. The dorsal coloration of adults of Masti- copbis f. flagellum ranges from a completely pale tan dorsum with narrow, dark brown crossbands to one almost completely black dorsally. The most widespread type of col- oration has been described above. There are disjunct populations in northern and central Florida and southern Georgia, the adults of which are characterized by a pale tan dorsum with narrow, dark brown crossbands. A specimen of this phase (LSUMZ 18527) that I collected on 30 March 1968 on St. George Island, Franklin County, Florida, was tan dorsally with very dark brown pigment on the posterior edges of the internasals and prefrontals, the pos- terior one-half of the frontal, most of the supraoculars except for the extreme an- terior portion, and the anterior and pos- terior portions of the parietals. There is some dark brown pigment on the neck arranged in the form of indistinct bands. The first band is the best developed and is about four scales wide. Ventral coloration is cream with a triangular extension of the dorsal ground color onto the lateral edges of the ventrals and some tan and dark mottling on the neck. The chin is white, heavily mottled with tan and dark brown pigment. The specimen is an adult male, measuring 1280 mm in snout-vent length. This color phase is known from the fol- lowing localities in Florida (Fig. 4): Ala- chua County, 1 mi. NE Archer (UF 9656); Clay County, 3 mi. S Middleburg ( UIMNH 25699), Gold Head State Park (TCWC 10781); Duval County, near Jacksonville (UMMZ 67807); Escambia County, Pensa- cola ( MCZ 98 ) ; Franklin County, St. George Island (LSUMZ 18527); Gulf County, 2 mi. S Wewahitchka (UF 6546); No. 2 Coach whip Snake, Masticophis flagellum 39 I \ f \ PICEUS FULIGINOSUS CINGULUM I* Figure 3. Distribution of Masticophis flagellum in Mexico. Explanation as for Figure 2. Jackson County, 1.5 mi. W Campbelkon (AU 2037); Lake County, Tavares (UF 113), Leesburg (UF 2456); Leon County, east of Tallahassee ( FSU 116); Liberty County, 6 mi. S, ^u> mi. E Bristol ( UF 10083-1, 10083-2); Marion County, Fort McCoy (AMNH 23100); Orange County, Orlando ( SDSNH 36740 ) ; Putnam County, road between Interlachen and Grandin (MCZ 67179); Washington County, 1 mi. N Ebro (LSUMZ 16860). In addition, Charles W. Myers (in litt.) informed me that two similar pale-phased specimens of flagelh/m escaped from him at the two following localities: Franklin County, 6 mi. S, 9V'; mi. W Carrabelle, near Green Point, and Liberty County, 18 mi. NNW Carra- belle on Florida Hwy. 67. The following records are available from Georgia: Baker County, near Newton, Pineland Plantation (CM 40196-7), 3 mi. N Newton (BYU 22188), Itchwa Plantation (UG 206); Baldwin County, Milledgeville (MCZ 286); Brantley County, 20 mi. SE Waycross (UMMZ 92793); Decatur County, 4 mi. SW Bainbridge ( INHS 7612 ) ; Long County, 10 mi. SE Thomasville (TCWC 11945), 1 mi. S Thomasville (TCWC 11943), 3 mi. NW Metcalf (TCWC 11944); Wilcox County, Benhill Fishery (UG 262). Charles W. Myers also informed me (in litt.) that he saw a DOR specimen of this phase 9^2, mi. N, 5 mi. W Bainbridge on U.S. Hwy. 27, Decatur County, Georgia. He stated that the specimen "was also uniformly light colored but probably not as pale as those from Florida ..." In addition, I have seen one specimen of this phase ( NCSM 1966) from Rose Hill, Duplin County, North Caro- lina. Even excluding this pale phase, the dorsal coloration of adults tends to be paler in Florida than in other portions of the range ( Fig. 5 ) ; the darkest animals occur in Mis- souri, northwestern Arkansas, and eastern Oklahoma (Fig. 6). In dealing with an ani- mal that has a color pattern consisting of an evenly graded change in color from one end of the body to the other it is difficult to devise an objective system for evaluating this color pattern. Variation in M. f. flagel- hnn, therefore, is more difficult to discuss 40 Tulane Studies in Zoology and Botany Vol. 16 Figure 4. Distribution of the pale phase of Masticophis flagellum flageUum in Florida and surrounding areas. than that of any other subspecies. In record- ing color notes on specimens, I have made an arbitrary judgment regarding the in- tensity of the color present and the extent to which the anterior portion of the body is covered by dark pigment. An estimate of the amount of dark pig- ment was made on the basis of a fraction of the total body length, i.e., 1/4, 1/3, 1/2, 2/3, or 3/4. In terms of this crude method the following statements can be made (Ta- ble 1 ) . Specimens from Florida are pre- Table 1 Percentage of specimens of Masticophis f. flagellum exhibiting various amounts of dark pigment on the anterior portion of the body. The amount is expressed as a fraction of the total length. Area Fraction of Body Covered by Dark Pigment 1/4 1/3 1/2 2/3 3/4 Florida 68 16 16 0 0 North and South Carolina 12 71 17 0 0 Georgia, Alabama, and Mississippi 26 39 30 0 5 Louisiana, Kentucky, Tennessee, and east- ern Texas 0 20 50 20 10 Arkansas, eastern Oklahoma, eastern Kansas, and Missouri 0 5 48 33 14 dominantly pale, usually one fourth or less of the body is covered with dark pigment. The extent of dark pigment on the body increases clinally from Southeast to north- west, those specimens from Arkansas, Okla- homa, Kansas, and Missouri being darkest. In addition, there is also a tendency for specimens from the southeastern and eastern portions of the range to be tan on the pos- terior portion of the body and for those from the northwestern and western portions of the range to be reddish brown to dark reddish brown on the posterior portions of the body (Table 2). The color of the pos- Figure 5. Masticophis flagellum flagellum from Sanibel Island, Lee County, Florida. No. 2 Coachu'hip Snake, Masticopbis jlagellmn 41 Fimue 6. Ma.sticophis jlagcllum flai^elltiiii from Salina Creek, 2 miles S Saliiia, Mayes County, Oklahoma. terior portion of the body normally ranges from tan through light brown, brown, and dark brown to black. I have seen only one specimen, however, that was black pos- teriorly (OSU 3244 from Okmulgee County, Oklahoma ) . Occasional specimens throughout the range of jlageUiim have a series of light bands across the body ( Fig. 7 ) . These bands range from four to ten scales in length and range in number from one to eight in any one snake. Some specimens ( especially sev- eral from Mississippi, Louisiana, North Table 2 Percentase of specimens of Masticophis f. jlagcllum exhiliiting various colorations of the posterior portion of the body. Light Dark Area Tan brown Brown brown Florida 91 9 0 0 Georgia 58 31 11 0 Alabama 41 41 18 0 Mississippi 83 13 4 0 South Carolina 80 20 0 0 Kentucky, Tennessee, and Illinois 75 25 0 0 Louisiana 47 16 35 2 Eastern Texas 19 33 31 17 .Arkansas 19 25 37 19 Eastern Oklahoma 2 28 42 28 Eastern Kansas 0 66 17 17 Missouri 10 19 52 19 Carolina, and eastern Texas) have very dis- tinct bands that are tan in color in contrast with the dark to very dark brown areas that surround them and extend completely across the dorsum. These bands may be either uni- colored or bisected transversely by a narrow dark brown band through the center. Other specimens show only slight evidence of this light banding, which is confined to the mid- dorsal and one or two paravertebral rows of dorsal scales. I have seen banded specimens from every state within the range of flagel- him except South Carolina, Kentucky, and Illinois (only one adult from each of the last two states). There is no geographical continuity to this type of pattern, inasmuch as specimens from the same general area may be banded or nonbanded. For example, in a series of twelve adults from Bladen, Hoke, Richmond, and Scotland counties in south-central North Carolina, eight speci- mens are banded and four are not. My interpretation of the zone of inter- gradation between M. f. flagellum and M. f. testaceus is at variance with that given by Brown { 1950). Examination of the material from eastern Texas demonstrates that the southern portion of the intergrade zone ( Fig. 2 ) is fairly wide, as indicated by Brown, but lies farther to the east. My rea- son for this conclusion is that specimens from Colorado County ( KU 1459, 1463), and Matagorda County (MCZ 19895, 42 T/dane Studies in Zoology and Botany Vol. 16 liH Figure 7. Pattern at inidbodv of Masticophi.s flaocUum flagclhmu (CM 24649) from 1.7 miles S Boles, Scott County, Arkansas. UIMNH 28508, USNM 32770-71 ) show no approach to flagelh/m in coloration. Also, I have seen three specimens froin Galveston Island, two of which (UIMNH 25811, UNM 8849) are typical test ace iis whereas the other (UIMNH 28510) shows some in- fluence from jlagellum. Specimens from Bell County, from which Brown listed one speci- men with no specific locality, are typical testacens ( KU 72914, SM 7009 and 7033, UNM 9207). Another specimen (KU 72915) from Bell County may show some influence from flagelh/m. On the eastern edge of the intergrade zone two specimens (TCWC 196, TNHC 33876) from Harris County are intergrades and two others (ANSP 10871, LSUMZ 14146) show no approach to testacejis. Most of the 14 speci- mens from Brazos County that I have examined are jlagellum, but five (TCWC 195, 2482, 4751, 14522; UAZ uncata- logued) show influence from testaceus. North of Dallas and Tarrant counties the zone of intergradation apparently becomes very narrow. I have seen a single specimen from Wise County (USNM 83127), most likely the same specimen examined by Brown, and I agree with him that it is a testaceus showing no approach to flagellum. I have been fortunate in having available a series of recently collected specimens from Denton County. All four adults from this county (NTSU 193, 238, 268; TNHC 26773 ) definitely have a flagellum pattern with no influence from testaceus. One juve- nile (NTSU 223) is unquestionably a //rf^^/- lum as it has already begun development of the typical adult pattern of this subspecies. Two other juvenile specimens (NTSU 186, 207 ) look like testaceus. Neither specimen, however, has the dark head markings of juvenile flagellum. I do not know if juve- niles of the two races can be distinguished in an area of intergradation. The zone of intergradation between flagel- lum and testaceus in Oklahoma and Kansas is largely conjectural. A single specimen ( KU 8389) from Love County, Oklahoma, is a flagellum. I have seen no specimens between '^his county and Payne County, Oklahoma, to the north. Most of the specimens I have seen from Payne County are flagellum, but four (MCZ 61175; OSU 482, 485-86) show influence from testaceus. The sample from Noble County to the north includes some of the darkest specimens of flagellum that I have examined. I have seen no intergrade specimens from Kansas, and the zone of intergradation that I depict (Fig. 2) is based on Smith (1950). The zone of intergradation follows ap- proximately the ecotone between the pine and hardwood forests of eastern Texas and the prairie and hardwood forests of east- central Texas ( Austroriparian and Texan biotic provinces of Raun, 1965). According to Raun the major trees of the Austrori- parian biotic province include longleaf pine (Pinus palustris), loblolly pine (Pinus taeda), yellow pine (Pinus echinata) , post oak (Quercus stellata), blackjack oak {Ouer- cus marilandica) , red oak (Quercus fal- cata) , sweetgum (Liquidambar styraciflua) , and wax myrtle (Myrica cerifera) . The major trees of the Texan biotic province are post oak, blackjack oak, and hickory ( Carya spp. ) , and mesquite ( Prosopis juli- flora), a recent invader. Masticophis flagellum testaceus (Say) Western Coachwhip Coluber testaceus Say, 1823: 48 Psammophis flavi-gularis Hallowell, 1852: 178 (type, ANSP 5388, Cross Timl^ers, Okla- lioma ) Mdsticopliis fhiviguhiris: Baird and Cirard, 1853: 99 (part) No. 2 Coachwhip Snake, Masticophis flagellum 43 HcrpctodnjUfi flagellifoniiis: Ciinther, 1858: 118 (part) Con/phodon testaceus: Ciinther. 1858: 108 Masticophis testaceus: Baird, 1859: 43 Masticophis flagcllifonnis testaceus: Ian, 1863: 65 Masticophis flagcUifonuis tcstacea: Jan, 1867: li\r. 20, pi. 6, fi.u. 2 Bascaniutii flageUifonne testaceum: Yarrow, 1875: 542 (part) Bascanium flagcllifoniie hicinctum Yarrow, 1883: 153 (type, USNM 11814, Old Fort Cobb, Te.xas ) Cohiber flagelliforniis testaceus: Carman, 1883: 43 (part) Bascanion fhigcllifonuis: Dunicril and Bocourt, 1886: 705 (part) Zamenis flagelliformis: Boulenger, 1893: 389 ( part ) Zamenis flavigularis: Ciinther, 1895: 120 (part) Bascanion fhigellifoinie: Dnf>;es, 1896: 481 Zamenis flagcUum jlageUum: Cope, 1900: 799 ( part ) Bascanion fhigelhim: Bailey, 1905: 46 Bascanion fhigelhim frenatuni: Bailey, 1913: 35 Cohiher fhigelhim fhigelhim: Stejneger and Barbour, 1917: 79 (part) Masticophis fhigelhim flavigularis: Ortenberger, 1923: 2 Coluber fhigelhim jueeus: Van Denbnrgh, 1924: 217 Holotype. — Lost. Originally in the collec- tion of the Academy of Natural Sciences of Philadelphia. Type locality: junction of Turkey Creek with the Arkansas River, 12 mi. W Pueblo, Pueblo County, Colorado (Maslin, 1953). Diagnosis. — A subspecies of Masticophis flagellum characterized by a light tan or light brown ( Fig. 8 ) to pinkish red dorsum with darker narrow crossbands on the neck and a double row of dark spots on the otherwise immaculate cream belly. Range. — South-central United States from western Kansas, extreme southwestern Ne- braska, and eastern Colorado, south through eastern New Mexico and all of Texas ex- cept for the eastern pine forest. In Mexico the western coachwhip occurs in eastern Coahuila, Nuevo Leon, Tamaulipas, eastern San Luis Potosi, and extreme northern Vera- cruz ( Figs. 2 and 3 ) . Description. — The following description is based on LSUMZ 18688, an adult female from 10 miles SE Colorado Springs on the grounds of Fountain Valley School, El Paso County, Colorado, collected 18 May 1967 by Robert M. Stabler. This specimen is from relatively near the type locality. Scutellation: supralabials 8-8, 4th and 5th entering the orbit; infralabials 11-10, four touching the anterior chin sliields, fifth the largest; loreal 1-1; preoculars 2-2; postoculars 2-2; tem- porals 6-6; scale reduction pattern 3 + 4(9) 3 + 4(121) 19 17 ^ 3 + 4(10) 3 + 4(116) 7 + 8(134) 7=7 + 8(138) 15 7 + 8(131) 13 7 + 8(141) 14 ^ 13; ventrals 197; tail incomplete. Head-body length 1159 mm. The dorsum is Dragon's-blood Red (color terminology based on Ridgway, 1912) on the nape grading to Light Coral Red pos- teriorly. The dark crossbands on the neck are near Liver Brown. The venter is Light Coral Red. The spots on the ventral surface of the neck are Dragon's-blood Red, the re- maining spots are yellowish cream. The top of the head is near Hay's Russet (reddish brown ) . The anterior edge of each dorsal head scale is lighter (pink). The side of the head is reddish brown and the temporals are outlined with pink. A cream spot is present on the upper preocular. The chin is white with mustard yellow and reddish orange blotching. Discussion. — Meristic and mensural data for this subspecies, based on 365 males and 313 females, may be summarized as follows: supralabials 7 to 9, infralabials 8 to 12, pre- oculars 2 or 3, postoculars 1 to 3, loreal 1 to 2, ventrals in males 173 to 204, ventrals in females 180 to 203, subcatidals in males 95 to 121, subcaudals in females 93 to 117, dorsal scale rows 19-17-12 to 19-17-15. Body length in males of all ages ranges from 263 to 1480 mm, in females 279 to 1270 mm. Total length in males of all ages from 375 to 1780 mm, in females 376 to 1655 mm. Tail length /total length ratios in males range from 0.195 to 0.272, in females 0.224 to 0.272. Maslin (1953) noted the presence of red individuals of Masticophis flagellum in the Arkansas River Valley of eastern Colorado. He stated that this phase occupied a distinct area in Colorado and demonstrated that the name Coluber testaceus Say, 1823, was ap- plicable to this form. Prior to this date, 44 Tulane Studies in Zoology and Botany Vol. 16 Figure 8. Masticophis flagellum testaceus (LSUMZ 12878) from Meade County State Lake and Park, Meade County, Kansas. however, Schmidt and Smith (1944) and Jameson and Flury (1949) had noted the presence of red specimens of Masticophis jlagellufn in Trans-Pecos Texas. Schmidt and Smith {lac. cit.) stated "the red racer is a familiar snake on the desert areas surround- ing the Chisos [Mountains]." Jameson and Flury {loc. cit.) also stated that "two of our specimens have a distinct reddish color an- teriorly and ventrally while another is dark red." They noted, however, that "other speci- mens show the more typical yellow-tan dor- sal color." Minton (1959) and Axtell (1959) likewise noted the presence of red- dish-colored whipsnakes in the Big Bend National Park and the Black Gap Wildlife Management Area in Brewster County, Texas. Recently, I became aware of a similar population of red testaceus in eastern New Mexico. I have seen specimens from ^4 mile N Logan (JRD 9518) and the vicinity of Tucumcari ( UAZ uncatalogued ) , Quay County, 10 miles NW Ramon (UNM 172) and 8 miles ENE Vaughn (JRD 9519), Guadalupe County, and about 65 miles NW Roswell, Lincoln County (NMSU 2213). I examined the University of Arizona speci- men just a few days after it was preserved. The dorsum was reddish orange with about ten narrow darker bands on the neck and forebody. The venter was cream with a light reddish orange overtone and a double row of indistinct reddish orange spots on the neck. The existence of these disjunct popula- tions ( Fig. 9 ) of reddish-colored individuals in eastern Colorado, eastern New Mexico, and western Texas, and the presence of nor- mally pigmented individuals in at least the Trans-Pecos populations, makes Maslin's (1953) arrangement untenable. The name, therefore, of the wide-ranging tan form (as well as the red phase of this subspecies) should be Masticophis flagellum testaceus ( Say ) . The factors responsible for the existence of these red whipsnakes in at least three dis- junct areas remain obscure. The shortgrass prairie of eastern Colorado and eastern New Mexico, and the desert scrub and barren flats of the Big Bend region would seem to have little in common. Thus, the occurence of the red color phase seems not to be cor- related with any vegetation type. Considering the disjunct nature of the range of the red populations, there is no alternative to recognizing them as represent- No. 2 Coachwhip Snake, Masticophis flagellum 45 Figure 9. Distribution of the phases of Masti- cophis flagellum testaceus in the United States. Strippled areas indicate the range of the red phase and the numbers within the areas out- lined by a dashed hne are the relative per- centage values of the three pattern types of A/. /. testaceus. ing a color phase of the subspecies testaceus. The exact limits of the range of this form are difficult to plot inasmuch as the red coloration fades in preservative to the tan of typical testaceus. Maslin (1959) listed specimens from Baca, El Paso, Fremont, Kiowa, Las Animas, Otero, and Prowers counties, all in southeastern Colorado. In addition, Maslin (in litt.) stated that they occur in Bent County. In New Mexico it is known from the three counties listed above. In Texas it is found in Presidio and Brewster counties (Fig. 9). In contrast to the pink or red coloration, the tan coloration occurs virtually through- out the range. A snake with this type of coloration was described by Hallowell (1852) as Psamtnophis flavi-gu/aris. The name flavigularis was used for the western coachwhip by Ortenburger (1928), who thought that Say's (1823) description of Coluber testaceus as a "pale sanguineous or testaceous" snake too uncertain to warrant the use of this name. Klauber ( 1942, p. 93, footnote) expressed a contrary opinion, however, when he stated, "1 cannot see by what line of reasoning the name testaceus is not applicable to this subspecies [the western coachwhip]. At the time the type specimen was collected (about July 18, 1820) the Long Expedition was at the foot of the Rocky Mountains, toward the head- waters of the Arkansas River. The descrip- tion, though brief, fits no other snake found in that region. It does not fit the prairie racer, usually referred to as C. f. flavigularis, and this snake does occur in that vicinity. I am therefore of the opinion that the prairie subspecies should be known as Coluber flagellum testaceus Say, 1823." Ortenburger (1928) noted three basic types of coloration in M. f. testaceus. These were: "( 1 ) a very light brown ground color with no indication of a pattern; (2) the same general light brown ground color with nu- merous narrow, darker crossbands ( two to three scales in width), which extend for a considerable portion of the body length; ( 3) few wide, dark crossbands (ten to fifteen scales in width ) on either a light or dark brown ground color." Ortenburger further noted (p. 102): "Variation in color and pattern is prob- ably greater in this form than in any other member of the genus. When the actual distribution of the types of pattern was studied, it was found that there is a defi- nite relation between the types of pattern and distribution. To the east throughout the region between the ranges of flagel- lum and flavigularis it was noticeable that there is a much greater number of speci- mens which possess the pattern com- posed of a few wide stripes. Toward the center of the range and south the uni- color light form is more numerous; to the west this again is gradually replaced by the form with many narrow stripes. Southeastern Texas seems to be the only region which does not hold to this type of distribution, as here one finds (if we can trust the locality records) a veritable mixture of all three forms. At present it does not seem that these forms should 46 Tulane Stitdies in Zoology and Botany Vol. 16 be considered as distinct, since there are no structural characters which will sup- port the color pattern differences, and it is believed that the locality records for southeastern Texas, if accurate enough, would show that these forms are found in distinct habitats. While lack of data at present precludes the possibility of deter- mining definitely what it is that actually causes these differences of color pattern, there can be no doubt that definite rela- tions of pattern to environment will be shown when the necessary data are avail- able." My data suggest a somewhat different in- terpretation. Contrary to what Ortenburger contended, the three color pattern phases of testaceus do not occupy discrete areas within the total range of the subspecies. Figures 9 and 10 show that the narrow-banded phase (A) is by far the most common in all areas within the range. The incidence of occur- rence of this pattern type ranges from 52% to 94%. It is highest in Kansas and lowest in Trans-Pecos Texas. The incidence of the unicolor phase (B) ranges from 6% to 34%. It is lowest in Kansas and highest in Trans-Pecos Texas. The incidence of the widebanded phase (C) ranges from 0% to 20%. No specimens of this phase were found in the material from Kansas, Ne- braska, Oklahoma, or the panhandle of Texas. The percentage of this phase is high- est in Coahuila and Nuevo Leon. There are no areas other than the ones mentioned above where all three of the phases fail to occur. All three phases were present in sev- eral collections from a single county. All three pattern phases undoubtedly do occur in the red phase of this subspecies also, how- ever, I have not seen unicolor specimens which were red at the time of examination. I have seen unicolor specimens from within the ranges of the red phase and these speci- mens may have been red in life. The following is a description of the color pattern of the narrow-banded phase (based on notes made on a living specimen, now in the private collection of Janalee P. Caldwell, collected 5.8 miles NW Cotulla on Hwy. 468, LaSalle County, Texas). The dorsum is tan with a slight pink wash on the nape. The neck bands are brown, 1 to IV- scales wide and separated by 2V-! to 3 scales. The venter is cream ( light yellow on Figure 10. Percent frequency of the three pattern type.s of Masticophis flagellitin testa- ceus in Mexico. neck) with a double row of light brown spots on the neck. The head is tan dorsally, the scales outlined with white. The lateral area of the head is also tan with light yellow blotching on some of the scales. The chin is white with light brown blotching. The unicolor phase is uniform light tan dorsally and cream ventrally with a double row of brown spots on the neck and fore- body. The following description is of the wide- banded phase (based on TNHC 30448 from 4 miles SSW Cuatro Cienegas, Coahuila, Mexico; Fig. 11). The dorsum is tan with about 6 brown bands, 12 to 14 scales in width. Each of these long brown bands has several narrower, darker bands within it. The throat is dark brown and there is a double row of brown spots extending pos- teriorly from this area. The posterior portion of the venter is cream. The chin is dark brown with cream spotting. I can find no basis for Klauber's (1942, p. 93) statement that "C. f. flavlgtdaris . . . tends to produce black (especially an- teriorly ) specimens in parts of New Mexico I II No. 2 Coachivhip Snake, Masticophis flagellum 47 Figure 11. Masiicophis flagellum testacctt.s (TNHC 30448) from 4 miles WSW Cuatro Cienegas, Coahuila, Mexico. and Oklahoma ..." Dr. Klauber stated {in litt.) that although he had no memory of these specimens, perhaps some note of this characteristic might have been made on his original data sheets that he sent me. Upon looking through his data I find no mention of any testaceus from Oklahoma having this coloration. Two specimens, how- ever, which may have been in the University of New Mexico collection ( the designation used for the specimens is "N.M.," no num- bers are given ) from Bernalillo County, New Mexico, are described as "grayish black" and "light brown, somewhat darker on the head." I have examined the entire University of New Mexico collection and have seen no specimens fitting such descriptions. Possibly fading may have taken place as it often does in snakes that have long been pre- served. Masticophis jlagelluni lineatulus Smith Lined Whipsnake Bascauium flagelliforme testaceus: Cope, 1886: 284 (part) Masticophis flagellum flavigttlaris: Ortenburger, 1928: 102 (part); Dunkle and Smith, 1937: 4; Gloyd and Smith, 1942: 233 Masticophis flagellum lineatulus Smith, 1941: 394, 1943: 446; Tavlor, 1944: 185, 1949: 170, 1952: 811; Smith and Tavlor, 1945: 95, 1950: 327; Brown, 1950: 168; Schmidt, 1953: 189; Stebbins, 1954: 495, 1966: 150; Conant, et al, 1956: 12, 1965: 18; Grant and Smith, 1959: 55; Cochran, 1961: 196; FowJie, 1965: 71 Holotype.—USNU 105292. Type local- ity: 11 miles N San Buena Ventura, Chihua- hua, Mexico.^ Diag}70sis. — A subspecies of Masticophis jldgelli/m characterized by a tan or light gray dorsum with each dorsal scale on the anterior portion of the body having a cen- tral longitudinal dark streak (Fig. 12). The posterior portion of the venter and the un- derside of the tail are salmon pink (this color does not fade in preservative ) . Range. — Extreme southwestern New Mex- ico and southeastern Arizona south through most of Chihuahua ( except the southwest- ern portion), southwestern Coahuila, eastern Durango, northern Zacatecas, and western San Luis Potosi (Figs. 2 and 3). Description of Holotype. — Scutellation : supralabials 8-(S, fourth and fifth entering the orbit; infralabials 9-10, four touching the anterior chin shields, fifth the largest; loreal 1-1; preoculars 2-2; postoculars 2-2; temporals 7-8; scale reduction pattern 3+4(11) 3 + 4(119) 19 17 3 + 4(13) 3+4(121) 7 + 8(120) 7 + 8(142) 15 ^ 14 ^ 6 + 7 (173) 13 12; ventrals 195; tail incomplete. Body length is 16 16 mm (Smith reported 1481 mm, ap- parently in error ) , the incomplete tail mea- sures 335 mm. Smith (1941) described the color pat- tern of the holotype as follows: "Head light yellowish brown, darker toward posterior sutures; sides of head light yellowish brown, with a lighter area in the preocular, loreal, nasal and rostral; a dark, roimded spot in center of nasal 1 1 find this spot to be in the rostral, not the nasal | ; supralabials white below a line alx^ut even with posterolateral border of seventh labial and middle of sub- ocular labials. "Dorsal ground color light yellowish brown, becoming more reddish toward mid- dle of body, posteriorly mostly .salmon red; all anterior dor.sal scales with a central longitudinal black streak, which becomes 1 Original description stated "south" ( Coch- ran, 1961, p. 196). 48 Tulane Studies in Zoology and Botany Vol. 16 more spot-like on scales in middle of body, I would USNM 4388 from Laguna, Valencia barely indicated on posterior scales, as the County, New Mexico. Both have striping black spots become less distinct, the red on the dorsal scales but lack red color of areas become more distinct, the posterior the posterior ventrals and the subcaudals. scales being mostly red (with a little black There is very little pattern variation in near tip), with a white (cream) base; dor- adults of this subspecies, in marked con- sal surface of the tail is even more strongly trast to all other subspecies of Masticopbis marked with salmon red. flagelhnn with the exception of M. f. n/d- "Posterior edge of mental and broad areas docki. The color pattern of the specimens near the sutures between the infralabials and I have examined conforms very closely to chin shields, black-marked brown; a double the description given by Smith (1941). He row of black spots beginning with anterior failed to mention, however, the coloration ventral scales; posteriorly these spots becom- of the nape, which is light yellow to yellow- ing mixed with red and soon mostly red ish-tan. This coloration occupies an area of and very little black; anterior ground color from six to twelve scales posterior to the of belly yellow, this color extending onto parietals. This area is distinctly set off from lower dorsal scale rows; toward middle of the lineate coloration which follows it. belly this color is largely replaced by salmon There is some slight variability in the an- red, and posteriorly the belly is entirely red, terior extent of the salmon red ventral with the double row of black spots faintly coloration and the extent of coverage of indicated; ventral surface of tail mostly red, individual ventrals. In some specimens the the bases of the scales lighter (cream)." posterior half of each ventral is salmon red Discussion. — Meristic and mensural data or pink and the anterior half is cream. In for this subspecies; based on 29 males and others the whole scute is salmon pink or 20 females, may be summarized as follows: red. The ventral, as well as the dorsal, col- supralabials 8 or 9, infralabials 9 to 12, pre- oration varies ontogenetically and is dis- oculars 2, postoculars 2, loreal 0 to 3, ven- cussed below. trals in males 183 to 199, ventrals in fe- Specimens from Zacatecas are not so males 185 to 197, subcaudals in males 99 to brightly colored as those from Durango, 109, subcaudals in females 95 to 103, dorsal Coahuila, and San Luis Potosi. All three of scale rows 19-17-12 to 19-17-13. Body the adult specimens I have seen (UMMZ length in males of all ages ranges from 249 123251-52; AMNH 82156) lack bright yel- to 1270 mm, in females 290 to 1281 mm. low coloration on the anterior ventrals, and Total length in males of all ages ranges the salmon pink coloration on the posterior from 332 to 1650 mm, in females 386 to portion of the body and underside of the I6l6 mm. Tail length /total length ratios tail is not nearly so bright or extensive in in males range from 0.213 to 0.262, in fe- distribution. The nape band, which is light males 0.207 to 0.260. yellow in more typical adult lineatulus, is I have examined the entire type series of yellowish tan in the Zacatecas specimens, this subspecies and find that some comment Conant (1965) also mentioned this in his is necessary. Seven of the specimens (USNM discussion of AMNH 82156, which he re- 14279, 104675, 104676, 105292, 1988, ported as the first record for the state of 14283, 46355) are typical lineatulus. Two Zacatecas. (USNM 12676, 26151), from Guanajuato The record for this subspecies from 6 and "Mexico," respectively, are juvenile miles S Artesia, Eddy County, New Mexico Masticopbis striolatus. Another specimen ( Gehlbach, 1956), obviously is an error, (USNM 8429) from "New Mexico," con- inasmuch as the area is inhabited by tes- sidered by Smith (1941) to be a possible taceus. intergrade between lineatulus and jlavigu- Juveniles of Masticopbis flagellum linea- laris (= testaceus) , is a juvenile and can- tulus are indistinguishable from those of not be allocated to subspecies. Smith ap- M. f. testaceus, that is, in both races the parently considered USNM 1989 from Co- dorsal pattern consists of narrow (less than bre Mines, New Mexico, as typical of the one scale long to one scale long), uniformly subspecies; I would identify it as an inter- dark bands separated by lighter interspaces grade between lineatulus and testaceus, as of variable length (usually 2 to 3 scales in No. 2 Coach whip Snake, Masticophis flagellum 49 length ) . This differs from the type of color the venter, except for the first few ventrals pattern seen in juveniles of the /'/i:£'//'.f group, (which are cream) is bright yellow. Each in which the bands are longer ( usually of the dorsal scales has a more or le.ss cen- about 3 scales long ) and are bounded an- trally located black stripe. The nape is dull teriorly and posteriorly by an irregular, nar- yellow. row dark line. The dorsal surface of the Intergradation between AI. /. lineatulus and head in juvenile lineatulus is relatively uni- M. f. testaceus in New Mexico takes place form in coloration (unlike the head pattern over a broad area from Valencia and Ber- of flagellut)!, and the other subspecies in the nalillo counties southwestward to southern testaceus group), and the anterior portion Catron County and southeastward to western of the venter is marked with a double row Otero County. The southern limits of this of brown spots. There is a nonbanded area intergrade zone are unknown because of a of variable length ( 9 to 11 scales long ) on lack of specimens from northwestern Chi- the nape. huahua. Within New Mexico, however, By the time the animal has reached a there is decreasing indication of the linea- snout-vent length of approximately 500 mm, ti/lus pattern to the north. Specimens from the adult pattern has begun to develop. The Hidalgo, Luna, and Dona Ana counties show unique subcaudal and posterior body color a great amount of influence from lineatulus is one of the first of the adult characteristics to the south. to appear (indications of this coloration are The following color pattern is of a living shown by AMNH 82154 which has a snout- specimen from Las Cruces, Dona Ana vent length of only 305 mm). The salmon County ( LSUMZ 10519). The dorsum is color develops first along the posterior edge brownish gray anteriorly, grading to brown of the subcaudals and posterior ventrals. posteriorly. A series of indistinct bands is Also, the yellow coloration of the anterior present on the neck and forebody. Each region of the venter develops at about 500 scale on the anterior portion of the body mm. has a black line extending through the cen- The lineate pattern of the dorsum of the ter. The lateral edges of the posterior body adults develops from the banded juvenile scales are yellow. There is also an indistinct pattern by an intensification of the dark broken line on scale row 2 on the posterior markings on the scales within the juvenile portion of the body. The base of the scales bands and by the progressive development in row 1 is tinged with orange. The nape of longitudinal stripes on the scales between has a distinct yellowish tinge. The venter is the bands. Therefore, as the animal ma- cream to white with a double row of dark tures the salmon color becomes brighter, brown spots that become diffuse on the covers more area on each scale, and pro- forebody and are gradually replaced by light gressively obscures the banded juvenile pat- orange. This coloration becomes more prom- tern, inent posteriorly, and on the posterior por- As the lineate pattern of the dorsum be- tion of the body and tail forms the posterior comes more well defined, the nape area border of each scale. The head is light becomes more noticeable by its lack of pat- brownish gray above with darker blotching tern. Also, it is dull yellow in contrast to on some head scales. The lateral area of the the brown ground color of the dorsum. head is similarly colored with light yellow At about 900 mm the dorsum of the pos- blotches on the posterior nasal, loreal, and terior region of the body is infused with upper preocular. The supralabials are white the same salmon color that is found on the except for the posterodorsal corner of each, posterior ventrals. This salmon color occu- which is grayish brown. The chin is white pies the posterior half of each scale, leaving with brown blotching, the anterior half tan in coloration. Some New Mexico specimens from Dona In adult animals (above 950 mm) each Ana County show no pink or orange color- ventral on the posterior portion of the body ing on the ventrals or subcaudals and so is almost completely covered with salmon approach testaceus in this respect. A single pink pigment. The posterior half of each specimen from Catron County ( UNM subcaudal is also salmon-colored, the an- 9312) has no pink on the subcaudals but terior half being white. The remainder of has the typical lineatulus striping on the 50 Tidcwe Studies in Zoology and Botany Vol. 16 dorsum. A specimen from Grant County Portal and Rodeo on the eastern side of the (UNM 8422) is similar but has a pink Chiricahua Mountains. These individuals do tinge on the posterior edge of the sub- not differ essentially from specimens seen caudals. A specimen from Otero County from Hidalgo County, New Mexico. A speci- (NMSU 1796) has only slight indications men from 5-8 miles S San Simon in the San of striping on the dorsum and is otherwise Simon Valley (UAZ uncatalogued ) is simi- like testaceus. A single specimen from Sierra lar to those from near Portal and Rodeo ex- County (UNM 435 ) has light dorsal streak- cept that it lacks the salmon subcaudal color, ing and some light pink coloration on the and the lines on the dorsal scales are not posterior ventrals and subcaudals. particularly dark. It does not, however, show Specimens from Socorro, Valencia, and any influence from piceus. I have seen speci- Bernalillo counties show a varying degree of mens from the Sulphur Springs Valley and influence from lineatulus. Some specimens one from the San Bernardino Valley which more closely resemble testaceus, others Hnea- show influence from lineatulus and a varying I tulus. From its description, the specimen amount of influence from piceus and cingu- (TNHC 4487) discussed by Gehlbach hnn. Two specimens (both uncatalogued (1965) appears to be an intergrade. He specimens in the UAZ collections) appear stated "the recent specimen, an adult male, to be intergrades between piceus and linea- 1070 mm in total length, has longitudinal tulus. A specimen from 28 miles NW Will- lines on the dorsal scales but retains an- cox, at the north end of the Sulphur Springs terior crossbands and lacks the diagnostic Valley (UAZ uncatalogued), shows more red subcaudal color." influence from lineatulus than from piceus. Specimens from El Paso County, Texas, The following description is of this speci- also show characteristics of both lineatulus men, seen soon after it had been preserved, and testaceus. I have seen two such speci- The dorsum is brown with a dark streak mens (KU 72910, UMMZ 117769) . Robert down the center of each scale. Posteriorly G. Webb stated {in litt.) "strangely, these streaks become less prominent. There the coachwhips in the El Paso area are is an indication of banding on the neck as hard to come by — the only one I have in piceus. The venter is white anteriorly seen, which was a DOR, and is now grading to cream. The posterior edge of the a skeleton in our collections, I would call ventrals of the posterior region of the body an intergrade; although the specimen was and the subcaudals are salmon pink, torn up badly, the posterior part of the The other specimen is from the Willcox body (dorsally and ventrally) had the char- Playa and shows more influence from piceus. acteristic magenta color of lineatulus. This Its color pattern (in preservative) consists DOR specimen is from Jueco Tanks, about of a brown dorsum ( more reddish brown on 25 miles east of El Paso." Also, Strecker the neck and forebody ) with a few of the (1915) gave the following information: dorsal scales having lineatulus-type striping. "Several coachwhips from El Paso County Neck bands are of piceus type. The venter combine the characters of the two western is cream anteriorly with a double row of subspecies, frenatum and piceus, having nar- dark brown spots grading to light rust on row crossbands in front and the underparts the posterior portion of the body and the bright pink in color." These words, written underside of the tail. The head has a twenty-six years before lineatulus was de- typical piceus coloration, scribed, give a fairly accurate and succinct A specimen from 4 miles S Dos Cabezas description of a lineatulus X testaceus in- (ASDM 655) looks very similar to typical tergrade. banded cingulum, but there is a slight indi- In Arizona the intergrade zone becomes cation of lineatulus influence in neck band- narrower and more complex as three sub- ing. Another specimen from 5.6 miles N species intergrade in southern Cochise, Bernardino ( AMNH 75130) has a linea- southern Pima, and Santa Cruz counties, tulus nape band and a piceus-type head pat- M. f. lineatulus from the southeast, M. f. tern. Banding of a piceus type is present cingulum from the south, and M. f. piceus on the neck but lineatulus striping is also from the north. Influence from lineatulus present. Lowe (1964: 20) mentioned the is seen in specimens from the vicinity of intrusion of typical Chihuahuan Desert No. 2 Coachwhip Snake, Masticophis flagellum 51 vegetation into parts of the San Simon and Sulphur Springs valleys and it is evidently through these avenues that the influence of lineatulns has spread into Cochise County. M. f. piceus and M. f. ciiig//l//m inter- grade along the area of contact between the elements of the Sonoran Desert and the Desert-Grassland ecotone (see Lowe, 1964). Intergrades have been examined from the Palo Alto Ranch, along the Tucson-Nogales Highway (U.S. 89) and from about Con- tinental south to Arivaca Junction. I have also seen intergrades from the eastern slope of the Huachuca Mountains, and specimens which look like cinguhmi, piceus, and cingu- liim X piceus intergrades from the region of Fort Huachuca. Masticophis flagellmn cingulum is known in Arizona from several localities in the oak woodland between the Patagonia and Pajarito mountains and occurs as far north in Santa Cruz County as Tubac ( SDSNH 32504). I have seen one specimen from Peria Blanca in the foothills of the Pajarito Mountains, also an oak woodland habitat. Another specimen was examined from 8 miles S Arivaca (UAZ uncatalogued ) , and I collected a DOR specimen 1 1 miles NNE Arivaca, Pima County, in mesquite grass- land. M. f. cingulum is also found in desert grassland near Sonoita between the Santa Rita and the Patagonia mountains. It is also found in the region between Palominas and Douglas. Masticophis flagellum piceus is an in- habitant of the Sonoran Desert in Arizona and its distribution conforms well to the outline of that vegetational complex as de- scribed by Shreve and Wiggins (1964), except that it occurs somewhat farther east and south. It also extends north into the Mohave Desert. The type locality of M. f. piceus ( black phase ) , Camp Grant, is not included in the Sonoran Desert, as outlined by Shreve and Wiggins. This snake may provide additional evidence for a zoological redefinition of the eastern edge of the So- noran Desert (Lowe, 1955), but it does not occur as far east as Lowe has placed the boundary. It is difficult to decide exactly where the zone of intergradation between testaceus and lineatulus in southwestern New Mexico should be drawn. I examined a series of specimens from Hidalgo and Cochise coun- ties ( three specimens ) , and three from Chihuahua and Coahuila side by side, and found that if they were arranged in linear order according to snout- vent length the gradation in color pattern was not even. The Mexican specimens showed more character- istics of the subspecies lineatulus. The New Mexico and Arizona specimens showed fewer characteristics of lineatulus and more influence from the subspecies testaceus. I consider, therefore, that popula- tions of Masticophis flagellum from the southwestern portion of New Mexico and extreme eastern Cochise County, Arizona, represent intergrades between lineatulus and testaceus, albeit much closer to lineatulus. Masticophis flagellum piceus (Cope) Red Racer, Western Black Racer Bascanium fla^cUiformc testacetim: Cope, 1875: 40 (part) Bascanium fla^cUiforim' piccinii Cope, 1875: 40 Bascanion flageUifoniiis: Dumeril and Bocourt, 1886: 704 Zamenis flageUifoniiis: Buulenger, 1893: 389 (part) Bascanion flagellum frcnatum Stejneger, 1893: 208 (type: USNM 16340, Mountain Spring, Colorado Desert, San Diego County, Cali- fornia ) Bascanion piccum: Van Denburgh, 1896: 1006 Zamenis flagellum flagellum: Cope, 1900: 789 ( part ) Zamenis flagellum piceus: Cope, 1900: 804 Bascanium flagellum: Meek, 1906: 15 (part) Zamenis flagellifonnis frenatus: Ditniars, 1907: 288 Zamenis flagelliformis piceus: Ditniars, 1907: 288 Coluber flagellum frenatus: Grinnell and Camp, 1917: 190 Coluber flagellum piceus: Stejneger and Bar- bour, 1917: 79 Masticophis piceus: Ortcnhurger, 1923: 2 Masticophis flagellum frenatus: Ortenlnirger, 1923: 2 Masticophis flagellum frcnatum: Klanber, 1926: 144 Masticophis flagellum piceus: Tanner, 1927: 57 Holotype.—\]S,NU 7891. Skin of an adult female ( ? ) from Camp Grant ( = Fort Grant), Graham County, Arizona, collected by E. Palmer. Diagnosis. — A subspecies of Masticophis flagellum with two color phases, one black and the other red. The former is completely black dorsally and salmon pink to red posteroventrally. The red phase is pink to red with relatively wide crossbands on the 52 Tidane Studies in Zoology and Botany Vol. 16 Figure 12. Pattern on the anterior portion of the body of Masticophis flagellum Uneatiilus (MSU 4448) from 3 miles E Las Nieves, Durango, Me.xico. neck that range in color from that of the general ground color ( in which case the anterior and posterior edges are outlined with darker pigment) to black (Fig. 13). Range. — Arizona south of the MogoUon and Coconino plateaus, excepting Santa Cruz County and southern and southeastern Cochise County, then north through Mohave County, Arizona, into Washington County, Utah. From there the range extends west- ward into southern and western Nevada and southern and southeastern California exclusive of the San Joaquin Valley. In Mexico this subspecies occurs only in the Gran Desierto of northwestern Sonora and that part of Baja California del Norte that lies east of the San Pedro Martir Mountains and north of Bahia San Felipe (Figs. 2 and 3). Description of Ho/otype. — Scutellation: supralabials 8-8, fourth and fifth entering the orbit; infralabials 9-10, four touching the anterior chin shields, fifth the largest; loreal 1-1; preoculars 2-2; postoculars 2-2; temporals 8-7; ventrals 194; caudals 106 (tip of tail broken off). The holotype of Bascaniun? piceum Cope is a representative of the black phase of this subspecies. Inasmuch as the colors of the living animal have been lost or subdued by its long preservation the following descrip- tion of a specimen without precise locality' data and purchased from a dealer is given. The dorsum is black except for a few small flecks of salmon pink on the frontal and parietals. The venter is black anteriorly be- coming increasingly salmon pink. The lateral Vi of each ventral is black. The distal end of the dorsum of the tail is salmon pink with black flecks. The head is black except for a white spot on the upper preocular, the first and fcnirth infralabials, and a few gulars. Discussion. — -Meristic and mensural data for this subspecies, based on 232 males and 145 females, may be summarized as follows: supralabials 7 to 9, infralabials 9 to 12, preoculars 2 or 3, postoculars 2 or 3, loreal 1 or 2, ventrals in males 183 to 205, ven- trals in females 184 to 205, subcaudals in males 97 to 120, subcaudals in females 96 to 115, dorsal scale rows 19-17-12 to 19-17-14. Body length in males of all ages ranges from 294 to 1290 mm, in fe- males 300 to 1154 mm. Total length in males of all ages ranges from 393 to 1668 mm, in females 400 to 1541 mm. Tail length/ total length ratios in males range from 0.221 to 0.326, in females 0.233 to 0.298. The more widespread red phase of the subspecies piceus was described by Stejneger as Bascanium flagellum jrenatum. The scu- tellation data for its holotype (USNM 16340) is: supralabials 8-8, fourth and fifth entering the orbit; infralabials 11-9, four touching the anterior chin shields, fifth the largest; loreal 1-1; preoculars 2-2; post- oculars 2-2; temporals 5-6; scale reduction pattern 3 + 4(10) 3 + 4(110) 19 17 3 + 4(11) 3 + 4(112) 7 + 8(112) 7 + 8(128) 15 14 13; ventrals 194; subcaudals 114. The coloration of the type specimen is faded, and therefore the following descrip- tion is given of a specimen (LSUMZ 10058) which I collected 9 miles WNW Westmorland, Imperial County, California, on 10 August 1965. The dorsum is reddish orange anteriorly grading to pinkish tan No. 2 Coachwhip Snake, Masticophis flagellum 53 Figure 13. Red phase of Masticophis flap.clhnn i>iceus from Miramar, San Diego County. Cali- fornia. posteriorly. The first two neck bands are somewhat more brown than the ground color. The venter is light orange anteriorly grading to bright pink posteriorly. There are a few darker flecks on the ventral sur- face of the neck. The head is grayish brown dorsally and laterally. There are light mark- ings present on the posterior nasal, loreal, preoculars, postoculars, and temporals. The temporals are tinged with pink. The chin is white with gray flecks. The most significant aspect of color pat- tern variation in this subspecies is the sym- patric occurrence, in certain parts of the range, of two dimorphic phases, a black and a red one. As can be seen above, these phases were originally described as distinct species and were retained at that level by Or- tenburger (1928). He was of the opinion that the black and red phases differed in at least four other characteristics besides color. He stated that the black phase snake "is noticeably a stouter and larger animal than frenat//s," that the red phase snakes take to the bushes as a means of escape, whereas black phase snakes escape into holes in the ground, that the subcaudals are more num- erous in black snakes than in red ones, and, finally, that black snakes have a propor- tionally longer tail than the red snakes. Smith (1941) pointed out that pice//s is "almost beyond question, a mutant form, oc- curring within the range of 'fremalus'," and that the name frenat/ts is not available, be- cause it is a junior secondary homonym of Coluber jrenatus ( = Elaphe jrenata ) and thereby permanently rejected. Klauber (1942), in an excellent paper on the subject, discussed the status of the black whipsnake. In this paper he showed that each of Ortenburger's alleged differ- ences between the black and red phases was untenable. He also pointed to the existence of intermediate specimens as indicative of the conspecificity of the two forms. Further- more, he stated that this is not the only ex- ample of color pattern variation in this sub- species, to wit "coastal San Diego specimens have black necks, while those from the des- ert side of the mountains and the Imperial Valley are without this dark color or it is only faintly evident." No one seems seriously to have questioned this interpretation, with the exception of Brattstrom and Warren (195.3). In their description of Masticophis jlagellmn rud- docki they stated (p. 179) "because we have described riiddocki primarily on the basis of color, we feel that it might be well to clarify our position on the taxonomic status of the black forms of M. f. piceus in Baja California and Arizona. No new additions have appeared to challenge Klauber's (1942) conclusions that the black racer {M. piceus) is merely a color phase of the red phase (M. f. jreuatum) ( = M. /. piceus). We feel that intensive study will support some of Ortenburger's (1928) observations on the distinct ecological .separation of these two forms. Ortenburger, however, came to the conclusion that the black racer (M. 54 Tulane Studies in Zoology and Botany Vol. 16 piceus) deserved specific recognition. We feel that because of the existence of a few specimens which appear to be intergrades, the black racer should only be considered a race of the red racer (M. /. piceus and M. f. jrena- tum, respectively ) . As for the black forms of southern Arizona, it is quite possible that these represent another subspecies whose closest affinities are with the black racers of Baja California. ( If so, it would become M. f. piceus and the Baja California form named new). Such a disjunct distribution need not cause undue surprise in light of other recent works dealing with the same situation in almost the same area. We refer to the relationship between Crotalus viridis helleri and Crotalus viridis cerberus (Klau- ber, 1949) and to the possible relationship of Lampropeltis zonata to L. doliata gentilis (Zweifel, 1952)." These authors seem to have misunderstood Klauber's paper spe- cifically and the subspecies concept gen- erally. Klauber rightly pointed out in his 1942 paper (p. 88) that "since there ap- pears to be no area in which the black form is found alone, it can hardly be considered a valid subspecies." Thus, Brattstrom and Warren's statement that the intermediate specimens support the recognition of the black and red racers as subspecies of the same species is meaningless. The rest of the paragraph also has little pertinence. The situation in the whipsnakes is not at all comparable with that of Crotalus viridis cerberus and C. v. helleri, or Lampropeltis zonata and L. triangulum. The distribution of the black phase of piceus (Fig. 14) is confined to south-central Arizona. I have seen specimens from as far north as 15 miles SE Florence, Pinal County, and Klauber (1942) reported a specimen from a little farther north at Florence Junc- tion. I have seen the phase as far west as 25 miles E Gila Bend, Maricopa County, and 5 miles N Park Headquarters, Organ Pipe Cactus National Monument, Pima County; as far south as Sasabe, Pima County, and the Santa Rita Experimental Range, Pima County; and as far east as Fort -Grant ( = Camp Grant, the type locality of piceus), Cochise County. In order to determine, at least approx- imately, the density of distribution of the black phase over its range I placed a 10- mile square grid over a standard highway Figure 14. Distribution of the black phase of Masticophis flagellum piceus in Arizona. map of Arizona and recorded the number of specimens that fell within each section of the grid. I also recorded the numbers of specimens of typical red phase piceus and those with intermediate coloration. Of the specimens of the black phase examined, 75% were collected within all the squares touching the square occupied by Tucson in Pima County. Beyond this area the percent- ages become smaller, although more black phase individuals occur west of Tucson than in other directions. The most distant record is about 110 air miles from Tucson, in the Organ Pipe Cacttis National Monument (UAZ uncatalogued). Other outlying rec- ords lie between 60 and 90 miles from Tucson. Klauber (1942) stated that black racers outnumber red ones in the vicinity of Tucson. In order to determine whether or not this is the case, I counted the numbers of the red, intermediate, and black snakes within each section of the grid from which specimens were available. My determinations made by this method corroborate those of Klauber. Within all the squares touching the square occupied by Tucson, where the black phase is most common, of the total number of Masticophis flagellum piceus ex- amined, 58.99 are of the black phase, 26.3% are of the intermediate phase and No. 2 Coach whip Snake, Masticnphis flagellum 55 14.8% are of the red phase. It is also in- teresting that although the black racer in- habits an area encompassing most of Pima and Pinal counties and portions of Maricopa and Graham counties, the snakes intermedi- ate in coloration between the black and red phases are only found within a 30-mile radius of Tucson. A specimen of this intermediate phase (LSUMZ 10521) from the vicinity of Tucson, Pima County, was given to me by John W. Wright. The neck and forebody of this specimen are black grading to reddish- brown, speckled with black markings on some scales. The middorsal area is more red than the lateral area. The venter is black on the neck grading to pink on the rest of the venter. The head is velvety black dorsally and laterally. The chin is very dark brown with a small amount of white blotching. The eye is gold around the pupil and dark brown around the edge. Unfortunately I have seen very few living specimens of the red phase of picens and comments on color are taken from the lit- erature. One feature of the color pattern that can be observed on preserved specimens is the difference in coloration of the neck bands. Klauber (1942) stated "coastal San Diego County specimens have black necks, while those from the desert side of the mountains and the Imperial Valley are with- out this dark color or it is only faintly evi- dent." The degree of variability over the entire range of picens is great. The neck bands may be the same color as the dorsum ( outlined anteriorly and posteriorly by a very narrow band of darker pigment) or range from light brown or light gray through grayish brown to black. I have sub- divided this continuous variation into five arbitrary categories: A. pink (essentially the same color as the dorsum), B. light gray or light brown, C. brown or gray, D. dark gray or dark brown, E. black. I have excluded those specimens having a coloration intermediate between the black and red phases. By this method a crude estimate can be made of the incidence of occurrence of each band color type in any given area ( Table 3 ) . With the exception of coastal southern Cali- fornia, which apparently lacks pink-banded individuals, all the neck band colors occur in every area, although one type may pre- dominate. Concerning the occurrence of Masticophis flagellum in the southwestern portion of Colorado, Maslin (1959) stated "the in- clusion of this species [meaning the sub- species piceus\ in the fauna of the state is based on reports of specimens seen in the vicinity of Mesa Verde National Park. I also saw, but failed to collect, a pale reddish-brown specimen 30 mi. W Cortez, Montezuma County, at the mouth of Yellow Jacket Canyon in June, 1958. It is possible that the form that undoubtedly occurs in this area should be assigned to AI. /. flavi- gularis. But biogeographic evidence, based on the distribution of other species of rep- tiles, indicates that it should be tentatively recognized as M. f. pice//s until corrobora- tion from actual specimens is possible." Douglas (1966), who spent three years at the Mesa Verde National Park, did not men- tion this subspecies in his paper on the am- phibians and reptiles of the park and, to my knowledge, no specimens have been col- lected to date. Douglas (pers. comm.) also stated that he had never heard of any red racers being found in this area. It seems un- likely that picetis or any subspecies of flagel- hnii would occur in this area since no speci- mens are known from the higher elevations of northeastern Arizona and northwestern New Mexico and, indeed, the distribution of Masticophis jlagellum seems to be limited by these higher elevations. Masticophis flagellum ruddocki Brattstrom and Warren San Joaquin Whipsnake Coluber flagellum jrenatus: Grinnell and Camp, 1917: 190 (part) Coluber flagellum piceus-.y/an Denburgh, 1922: 664 (part); Klauber, 1942: 88 (part) Masticophis flagellum frenatus: Ortenburgcr, 1928: 112 (part) Masticophis flagellum ruddocki Brattstrom and Warren, 1953: 177; Cunningham, 1955: 218; Conant, et ai, 1956: 12; Miller and Stebbins, 1964: 409; Montanucci, 1965: 278; Stelilnns, 1966: 150 Masticophis flagellum i)iccus: Stebbins. 1954: 495 (part) Holotype. — UCLA 6108 (specimen now in the collection of the Museum of Verte- brate Zoology, University of California at Berkeley), collected 't mile S of ridge at Wheeler Ridge and 1^-j miles W U.S. Hwy. 99, Kern County, California, on 12 April 1953 by James W. Warren. 56 Tulane Studies in Zoology and Botany Vol. 16 Table 3. Percentage of specimens of the red phase of Masticophis flagellum piceus exhibiting various colorations of the neck bands. Color of neck bands Light gray or Brown or Dark gray or Area Pink light brown gray dark brown Black Arizona 30 25 11 9 25 Utah and Nevada 11 20 33 13 23 Desert areas of southern California 15 8 8 15 54 Coastal areas of southern California 0 3 6 8 83 Diagnosis. — A subspecies of Masticophis flagellum characterized by a light yellow to olive yellow dorsum with neck bands obso- lete or absent ( Fig. 15 ) . Range. — The San Joaquin and Sacramento valleys of central California from Colusa County south to Kern and San Luis Obispo counties ( Fig. 2 ) . Description of the Holotype. — (based on data from original description ) . Scutellation : supralabials 8-8; infralabials 10-10; preocu- lars 1-1; postoculars 2-2; ventrals 197; sub- caudals 105. Brattstrom and Warren (1953) gave the following description of the color pattern (presumably in life). "The color varies from spots of Aniline Yellow on the sides of the head and neck to a solid ground color of Buffy Citrine on the sides of the body and Saccardo's Olive dorsally. The ventral color is consistently Straw Yellow (capitals are Ridgway's Color Standards, 1912), with Aniline Yellow or black on the lateral edges. The top of the head is light brown with two dark spots on the supraoculars. There is a small postocular brown stripe on the edge of the postocular and anterior tem- porals. The labials each have a dark spot and the anterior ventrals have Aniline Yel- low spots on them. There are two faint lat- eral light stripes on the anterior one-third of the body. This is caused by the first scale row being yellow-tipped at its dorsal and ventral corners. There is no evidence of any neck banding." Discussion. — Meristic and mensural data for this subspecies, based on 10 males and 6 females, may be summarized as follows: supralabials 8 or 9, infralabials 9 to 11, pre- oculars 2 to 4, postoculars 2, loreal 0 or 1, ventrals in males 188 to 198, ventrals in females 191 to 198, subcaudals in males 101 to 112, subcaudals in females 102 to 108, dorsal scale rows 19-17-13. Body length in males of all ages ranges from 764 to 970 mm, in females 987 mm. Total length in males of all ages ranges from 1015 to 1323 mm, in females 1329 mm. Tail length/ total length ratios in males range from 0.247 to 0.267, in females 0.257. I have seen only 17 specimens of rud- docki, excluding the holotype, the identity of which is made unquestionably clear by Brattstrom and Warren (1953). This subspecies, in common with M. f. lineatithts, exhibits none of the types of color pattern variation seen in the other subspecies and it has only one basic pattern. All of the specimens I have examined ex- hibit the coloration described for the sub- species by Brattstrom and Warren (1953), but there does seem to be some slight color variation. The dorsum ranges from yellowish tan to light olive yellow. Most specimens have at least some evidence of banding on the neck, but the bands are reduced in num- ber, in comparison with M. f. piceus, or are indistinct and, in general, the same color as the dorsum. Some specimens are light tan ventrolaterally ( to the level of scale row 5 or 6 ) and light olive yellow on the more dorsal rows. Several specimens ex- amined have a distinct orange tinge on the neck region, both dorsally and ventrally. The single specimen (SDSNH 26084) I have seen from the northern portion of the range ( actually in the Sacramento Valley ) is somewhat peculiar in having each dorsal scale with a darker central area giving the impression of a vague stripe on each dorsal scale row. No. 2 Coach whip Snake, Masticophis flagellum 57 •^ '■m- . .' ,» Figure 15. Masticophis flagellum nuldocki (UMMZ 78242) from 50 miles W Maricopa, Santa Barbara County, California. Apparently there is some color variation in this subspecies which reflects the influ- ence of Al. f. piceus. Joseph F. Copp ( /;; //■//.) stated that "... a DOR adult that was partly picked clean by birds was noted on California 166, 14.6 mi. W of New Cuyama, and was red rather than yellow as M. f. ruddocki is supposed to be. When they were fresh, the two specimens in my collection from that valley [JFC 62-62 and 63-55] were also red, and the black banding on the neck is moderately evident." Steb- bins (1966, pi. 30) illustrated a specimen of a coachwhip from Santa Cruz County, California, which is red. The specimen shows some neck banding but of a kind atypical for either ruddocki or piceus. The illustra- tion closely resembles the pink phase of M. f. testaceus. Intergradation with Masticophis f. piceus takes place in low passes through the moun- tains surrounding the southern end of the San Joaquin Valley. M. f. piceus occurs in Mohave, Great Basin, and Sonoran Desert vegetation in southern and southeastern Cali- fornia, extending as far north into chaparral as Los Angeles County to the west and Inyo County to the east. Intergradation may be expected anywhere along the southern end of the valley where there are areas in the mountains of sufficiently low elevation to allow contact between the two subspecies. I have seen an intergrade from 15.9 miles W New Cuyama, Cuyama Valley, San Luis Obispo County (JFC 63-55 ). Another speci- men (LACM 2239) from 5 miles W Ar- win cutoff, 3 miles S Bina Siding, Hwy. 466, Kern County, only about 20 miles from the type locality and within the valley, shows influence of piceus. It is yellowish tan above with gray neck bands, which are better developed than in mddocki. Five specimens were considered by Brattstrom and Warren (1953) to be intergrades be- tween ruddocki and piceus, and the localities for them are as follows: Kern River at Bod- fish (MVZ 2792), 6 mi. SSE Weldon (MVZ 56717), Weldon (MVZ 2793), Isa- bella (MVZ 2794), 1 mile E Onyx (UCLA 6106 = LACM 19258), and Monolith (LMK 27422 = SDSNH 27422). All lo- calities are in Kern County, four in the Kern River- Walker Pass area (5245 ft elevation), and one in the Tehachapi Pass area (3793 ft elevation ) . I have seen two of these speci- mens (LACM 19258 and SDSNH 27422) and consider both of them to be piceus with no evidence of the influence of ruddocki. Thus, the zone of intergradation between piceus and ruddocki as pictured in Fig. 2 is largely conjectural and roughly corresponds with the location of the mountains at the southern end of the San Joaquin Valley. The areas of intergradation between rud- docki and piceus need to be determined and 58 Tiihnie Stiaiies in Zoology and Botctny Vol. 16 the distribution of vuddocki within the val- ley needs to be better documented. Masticophis jlagellnm cingulum Lowe and Woodin Banded Red Racer Basccinion fhigcllum frenaUim: Van Denburgh, 1897: 463 Coluber flagcUum piceiis: Van Denburgh, 1922: 664 (part); Klauber, 1942: 88 (part); Bo- gert and Oliver, 1945: 363 Masticophis flagelhtin frcnatus: Ortenburger, 1928: 112 (part) Masticophis piceus: Allen, 1933: 11; Tavlor, 1936: 490 Masticophis flageUum frcnatum: Tavlor, 1936: 490 Masticophis jlagellnm frcnatum X flavigularis: Gloyd, 1937: 117 Masticophis flagelhnu piceus: Smith, 1941: 397 (part); Smith and Taylor, 1945: 95 (part); Langebartel and Smith, 1954: 134; Zweifel and Norris, 1955: 252; Stebbins, 1966: 150 (part) Masticophis flagellum cingulum Lowe and Woodin, 1954: 247; Conant, et al, 1956: 12; Fowlie, 1965: 69; Stebbins, 1966: 150 Holotype. — UAZ 672 collected at Mocte- zuma, Sonora, Mexico, about 2000 ft eleva- tion, in 1951 by A. W. Ruff. Diagnosis. — A subspecies of Masticophis flagellum with a highly variable color pat- tern. The dorsal coloration ranges from a series of wide reddish brown bands sep- arated by narrower, paired light bands to uniform reddish brown or uniform black. Range. — Arizona, from about midway be- tween Tucson and Nogales, east to Douglas, and west to the region of Arivaca, thence southward to southern Sonora, including most of that state except for the Gran Desierto and the mountainous eastern por- tion ( Figs. 2 and 3 ) . Description of the holotype. — Scutella- tion: supralabials 8-8, fourth and fifth en- tering the orbit; infralabials 11-12, four touching the anterior chin shields on the left and five on the right, fifth on the left and sixth on the right the largest; loreal 1-1; preoculars 2-2; postoculars 2-2; tem- porals 7-7; scale reduction pattern 4 + 5(11) 8 + 9(113) 19 — — 17 ^ 3 + 4(14) 3 + 4(117) 7 + 8(138) 16 ) 14 ^ 13; 3 + 4 117) ventrals 191; tail incomplete. Body length is 1023 mm, tail incomplete. Lowe and Woodin (1954) described the color pattern (in life) as follows: "Mid- dorsally the dark brown ground color tends to be blackish brown, grading to dark red- dish brown (near PI. 7, J 12)^ dorso- laterally and to lighter reddish brown (near PI. 7, J 7 ) laterally. Each dorsal scale tends to be darker on its posterior portion. On the anterior body, only a few scales are edged with pink (on the anterior portion of the scale); there is a progressive increase pos- teriad in amount of pink per scale. Thus pink predominates on the posterior body and tail and the darker brown is there restricted to the posterior edge of each scale. "The upper surfaces of the head are red- dish brown, distinctly darker on the supra- oculars and with faint light edges to the scales. There are conspicuous pink, ( PI. 2, J 9) marks (light areas) on the labials, postoculars, preoculars, nasals, and loreal. The pink of the postnasal, loreal, and pre- ocular is continuous and forms a "loreal stripe' from orbit to naris. Under surfaces of the head are variously colored cream and pink with small dark brown and black spots and blotches. "The ventral surface of the neck is dark pink (PI. 2, J 9 ) being approximately the same color as the undersurface of the tail and of the gular area. The ventral surface at mid-body is a lighter pink (PI. 2, G 8) than the neck and tail venters. The dark pink venter of the tail is near PI. 2, J 8. The entire ventral surface is patterned by bold brown transverse cross lines adhering to the posterior margins of the ventral scutes and extending completely across them. These are commonly associated with brown blotches on the lateral margins of the scutes. In ad- dition to the dark transverse lines on the neck, there are the commonly observed (in M. flagelhim) bilateral row of dark blotches and spots on neck and gular region gradu- ally fading away posteriorly. "On the upper surfaces, there is a series of conspicuous light-colored transverse, nar- row bands or cross-bars which extend com- pletely across the lateral and dorsal surfaces of the neck and body. Their effect is to 1 Their color terminology based on Maerz and Paul (1950). No. 2 Coachwhip Snake, Masticophis flagellum 59 break up the dark ground color of the 303 to 1-410 mm, in females 282 to 1186 upper surfaces into several large, longitu- mm. Total length in males of all ages ranges dinally oblong sections. The first of these from 408 to 1850 mm, in females 371 to bands is approximately four scales posterior 1541 mm. Tail length/total length ratios to the parietal head plates (counting scales in males range from 0.223 to 0.269, in fe- along the vertebral line). This anterior-most males 0.226 to 0.280. band is the narrowest and most sharply de- This subspecies was envisioned by its fined and is approximately one and a half describers as ranging from Moctezuma, So- scales in width. The second pink band is onora ( the type locality ) , northward to somewhat wider, being approximately two Amado, Santa Cruz County, Arizona. They scales in width, and, like the remainder, is stated, however, that the southern, eastern, less sharply delimited along its edges than and western limits were unknown at that is the first. The second band is approxi- time. The subspecies was diagnosed as fol- mately 12 scales posterior to the first. The lows: "characterized by adult dark red-brown third, fourth, and fifth bands are simple ground color on upper surfaces periodically single transverse bands approximately like broken by complete transverse, narrow, the second. The next and last three bands light-colored (pink) cross-bands which or band areas ( sixth, seventh, and eighth ) break the ground color into large, dark, are actually narrow double bands of pink longitudinally oblong sections; a single out- separated by a similar narrow band of dark standing light band crossing the nape." The brown ground color. The sixth band area approximate distribution of whipsnakes that (double band of pink) begins approxi- fit this concept of the subspecies cinguhim mately 75 scales posterior to the head (pa- is indicated by Stebbins (1966, map 132). rietals). It is separated from the seventh An examination of a large amount of ma- band area by approximately 18 scales; the terial from Arizona and Sonora indicates a seventh is similarly separated from the considerably more complex situation than eighth by approximately 17 scales. There was formerly recognized and requires a re- appears to be a faint trace of a ninth band definition of this form. This new informa- area on the posterior body half where the tion reveals that M. f. cmgulmn is distrib- ground color gradually becomes the uniform uted from Moctezuma in eastern Sonora to, braided pattern of the tail. and including, Isla Tiburon to the west. It "There is a faint, barely perceptible trace extends northward to intergrade with M. f. of a light longitudinal . . . line ( pink ) on piceus and M. f. lineat/dus in Pima and the anterior body involving the first two Cochise counties in Arizona. It occurs at scale rows. On the posterior body area, the least as far south as the Sonora-Sinaloa bor- lower scale row gradually becomes predomi- der. The situation in northern Sinaloa is nately pink, taking on the color of the ad- discussed in another section, jacent scutes. This is concomitant with the Masticophis jlagelhmi cingulum exhibits graduation posteriorly into a 'uniform' more pattern variation than any other race braided whiplike pattern of light pink and ^f flagellum. This has been the primary rea- brown. On the neck are traces of additional ^^^ for the confusion concerning Mastico- pmk 'bands' which give a clue to a probably ./^ -j. fiagelhmi in Sonora. Bogert and Oliver more banded neck m hatchlmgs and juve- ^ ^,^^^^ discussed the problem in part. They niles as is commonly seen in members of the , , , • • i r a i j ,, ,, , .. stated that their material from Alamos and jlapellum complex. „ . , • , i r u i V^. . A* • ■ J 1 J Guirocoba might be referrable to any one Discussjoti. — Menstic and mensural data , , i • . • for this subspecies, based on 84 males and ?."^ «V ' ^1'^?^^'^^^' t^"'"!' ^'f. '''!''.''' 41 females, may be summarized as follows: ^^'^^'^^^^^^^^^ and they noted that this being supralabials 7 to 9, infralabials 9 to 12, pre- ^^e case, "it becomes increasingly difficult oculars 1 to 3, postoculars 2, loreal 0 or 1, to allot specimens from critical areas, and ventrals in males 189 to 203, ventrals in for the most part, identifications are not females 188 to 201, subcaudals in males 97 based on characters but on geographical to 121, subcaudals in females 97 to 111, data." Taylor (1936) and Zweifel and Nor- dorsal scale rows 19-17-12 to 19-17-13. ris ( 1955 ) noted the considerable variability Body length in males of all ages ranges from of color pattern in the material they ex- 60 Tidane Studies in Zoology and Botany Vol. 16 amined, while continuing to use the name piceus. I suggest the following explanation for this situation. Masticophis jlagellum cingu- lum exhibits an unusual amount of variation in color pattern including a red phase (the phase originally described), a black phase (as in M. f. piceus and the Baja California form), a patternless phase (the phase dis- cussed by Bogert and Oliver, 1945, p. 363, and thought to resemble AI. /. testaceus), and intermediate patterns between these three phases. For purposes of discussion I have divided the wide spectrum of color pattern variation into seven pattern classes labelled as follows: A. Pattern of wide, dark red bands sep- arated by light pink interspaces; B. As in A but with a reduced number of light bands; C. As in A but with only the nuchal band present; D. Uniform pink or tan dorsal coloration with no evidence of pattern; E. As in A but with a few black scales distributed on the anterior portion of the body; F. Anterior 1/3 to 1/2 of the body black, posterior 2/3 to 1/2 red; G. Solid black dorsal coloration. The distribution of these color pattern classes is illustrated in Fig. 16. It will be noted that the individuals constituting the pattern A class tend to be concentrated in the northern and northeastern portions of the range. The patternless D class tends to be concentrated in southern Sonora, and the predominately black F and G classes are concentrated in central and western Sonora. The intermediate classes are rather indis- criminately distributed throughout the range. Figure 16 only approximates the actual distribution. The phases A, D, F, and G tend to predominate and there are a lesser number of the intermediate phases B, C, and E. In the samples examined, the incidence of the seven color pattern classes is as fol- lows: A— 35.6%; B— 5.9%; C— 2.5%; D— 12.7%; E— 12.7%; F— 23.7%; G— 6.8%. Also, as mentioned above, classes A, D, F, and G tend to be concentrated in cer- tain areas and also to predominate in those areas. In Santa Cruz County, Arizona, 84% are of pattern class A and 16% are of pat- PlCfUS C A ^V EO \EBa Figure 16. Distribution of the color pattern classes of Masticophis jlagellum cingulum in Sonora. tern class B. In Cochise County, Arizona, 89% are of pattern class A and 11% of pattern class D. From Naco, Sonora, south to Moctezuma, and west to Arizipe, 89% are of pattern class A and 11% of pattern class B. In the area from Imuris and Santa Ana south to Hermosillo, east to Mazatan and west to Pitiquito the following percentages obtain: A— 18%, B— 5%, C— 3%, D— 0%, E— 26%., F— 38%., G— 10%. From Desem- boque south along the coast to Empalme, including Isla Tiburon, the frequency of oc- currence of the various pattern classes is as follows: A— 8%, B— 4%, C— 0%, D— 12%, E— 15%, F— 46%, G— 15%. Finally, in the area between Cuidad Obre- gon, Alamos, and the Sonora-Sinaloa state border, 77% are of pattern class D, 15% are of pattern class C, and 8% are of pattern class B. Thus, it may be seen that the percentage of pattern class A is highest in the region from Santa Cruz and Cochise counties south to Moctezuma (90% of the total number of pattern class A ) . Percentages of pattern classes F and G are highest in the area from Santa Ana and Imuris south to Hermosillo and west to Guaymas and Desemboque ( 100% of the total number of pattern F and G classes ) . Finally, percentages of pat- tern class D are highest in the area from Obregon south to the Sonora-Sinaloa border No. 2 Coachivhip Snake, Masticophis flagellum 61 Figure 17. Pattern variation in Masticophis flagelhnn cinguliim. A. M. f. cin^uluni (hSVWZ 10520) from Douglas, Cochise County, Arizona (Pattern A). B. M. f. cinguhim ( LACM 6996) from Terreros, Sinaloa, Mexico (Pattern D). C. M. f. cingulum (AMNH 99370) from 0.5 miles S San Miguel de Horcasitas, Sonora, Mexico (Pattern F). D. M. f. vinculum (LACM 25922) from 25 miles N Hermosillo, Sonora, Mexico (Pattern G). E. A/. /. cingulum ( KU 78957) from 7 kilometers W Kino, Sonora, Mexico. 62 Tulane Studies in Zoology and Botany Vol. 16 (77% of the total number of pattern D class ) . There is variability in each of the above color pattern classes and, in reality, there is a complete continuum from the A pattern to the D pattern and from the A pattern to the G pattern. The basic coloration of pattern A is given in the following descrip- tion (based on color notes taken in life on LSUMZ 10520 from Douglas, Cochise County, Arizona; Fig. 17A). The dorsum has long, dark mahogany-red blotches sep- arated by narrower interspaces that are com- posed of scales that are light at the base and dark brownish red at the tip. Within each light interspace is a band that is some- what browner than the long bands. There are 10 or 11 dark red bands. These bands range from about 11 to 14 scales long. The light pink interspaces are 5 to 6 scales long and the dark band within these interspaces is about 2 scales long. Posteriorly, the long bands become less prominent and on the tail the color is like that of the interspaces on the more anterior portion of the body. Three scales behind the parietal there is a narrow (one scale long) band that is col- ored like the interspaces. The venter is dark pink with a slight orange cast. On the neck and forebody there is a double row of faint spots. The top of the head is dark red with the scales lightly outlined with white. The temporal scales are dark red. Most of the rostral is dirty white. The chin is white, blotched with tan and speckled with dark brown. The iris is golden orange and dark brown. The long dark bands vary from 5 to 16 scales in length and the light interspaces from 2 to 6. The number of dark bands on the body varies from 9-12. Frequently there is a narrow light band in the center of the dark band and occasionally two of these are present. I have seen no living material of pattern class B animals but the pattern, and prob- ably the color, is similar to that of pattern class A animals except that the long bands are reduced in number. Specimens that have a class C pattern are uniform pinkish tan, and the narrow light nuchal band is the only vestige of the class A pattern that remains. I also have seen no living specimens of pattern class D, but Bogert and Oliver (1945) described a specimen from Alamos as "uniform brownish olive above and es- sentially devoid of markings, although each of the lateral scales has a lighter area in the anterior median part. This specimen is simi- lar in coloration to specimens of Coluber flagellum testaceus from Texas, although its coloration is somewhat darker than any specimens at hand. The sides of the head, including the lips are almost uniformly brownish and bear no trace of pattern found commonly on piceus." These authors did not state whether the color description was based on a living specimen. Joseph F. Copp has informed me {in litt.) that "the species [Masticophis flagellum^ is also common on the same highway [Mexico 15] south of Navajoa where nearly uniformly red indi- viduals seem to be the rule." Preserved speci- mens are uniform tan to light velvety brown without any trace of bands (Fig. 17B). The venter is cream with a double row of light- centered spots on the neck and forebody. The top and sides of the head are the same color as the dorsum of the body. The lower portion of the supralabials is cream and they are often blotched with dark brown. The chin is cream, mottled with brown. The animals of pattern class D do, in- deed, appear similar to some specimens of Masticophis flagellum testaceus, especially the pattern variety that is devoid of any dor- sal markings. The most common pattern variant of testaceus is tan on the dorsum with narrow brown to dark brown bands on the neck, but the uniformly colored Sonoran specimens may be easily distinguished from this variety as well as from the wide-banded variety of testaceus discussed in the account of that subspecies. Separating nonpatterned cingulum and nonpatterned testaceus in- involves rather subtle distinctions, and, were it not for the fact that a continuum can be demonstrated from pattern A to D, it would be tempting to consider the convenient but zoogeographically untenable idea that non- patterned cingulum are testaceus. The vel- vety appearance of the dorsal coloration in nonpatterned cingulum is something that I have not seen in testaceus. Also the dorsal color seems to be darker in cingulum than in testaceus. Any attempt to delineate these differences in a key would be fruitless, and identification should be made on the basis of geographic probability. No. 2 Coach whip Snake, Masticophis flagellum 63 From the above it is obvious that a grad- ual obHteration of the banded pattern of class A animals occurs through classes B and C to pattern class D animals. This change is also indicated in the clinal shift- ing of the relative percentages of pattern A animals and pattern D animals from the northern portion of the range of cingidum to the southern portion. This same trend is also indicated to a lesser extent (due, perhaps, to lack of sufficient material) by the intermediate patterns B and C. Pattern E animals exhibit considerable variation and grade almost imperceptibly from the typical A pattern to the F pattern. At one end of the spectrum are animals similar to pattern class A animals but with 3 or 4 dark scales on the neck and some dark markings on the side of the head (KU 48921, JFC 63-117). Others have a few more black scales on the neck (LACM 9136, 25166; AMNH 75129, 84979-80; UAZ 9336; KU 78957, 78959; see Fig. 17E). Others have aggregations of black or very dark brown scales on the neck (JFC 64- 200 ) and farther posteriorly along the body (KU 78956, JFC 64-202, UIMNH 23917). This pattern type occurs primarily between Imuris and Guaymas. Pattern F animals also exhibit variability in the amount of the body covered by black (Fig. 17C). This black coloration is present on about 1/3 to 3/4 of the anterior portion of the body. The remaining 2/3 to 1/4 of the body is colored with a variegated mix- ture of red and black that, in many speci- mens, takes a banded form. These bands are very similar, and in some specimens identical, to the bands of pattern A animals except that the dark red pigment is re- placed by black. This type of banding is well evidenced in UNM 13705, MSU 7875, JFC 64-201, OSU 2029, and several un- catalogued specimens in the collection of the University of Arizona. More commonly, however, the black coloration is broken into long bands by lighter interspaces that are not bisected by a narrow black band. In these specimens the number of light bands varies from 2 to 7. Pattern G animals are solid black dorsally (Fig. 17D), although certain specimens have some dark brown pigment on the pos- terior portion of the body. One specimen (LACM 25922; Fig. 17D), which is black on the anterior 3/4 of the body grading to dark brown posteriorly with remnants of 3 lighter bands, is intermediate in colora- tion between those specimens that are classed as belonging to patterns F and G. The following description is of a living specimen of pattern class G (JFC 65-217). The dorsum is black except for the extreme tip of the tail which is blotched with pink. The venter is black on the anterior half of the body grading into pink on the posterior 1/4. The tail is dark pink ventrally. The head is very dark brown dorsally and laterally, except for a pink blotch on the loreal and upper preocular. The chin is very dark brown with fine white stippling on the anterior infralabials. The eye is golden orange around the pupil and dark brown around the edge. The pattern described above is identical with the pattern exhibited by black piceus from the region of Tucson. The two forms do not come into contact, however, and each is connected, through a pattern gradi- ent, with the more easily identified individ- uals of the subspecies cingulum and piceus. As is the case among the patternless phases of cin gill inn and testaceus, subspecific allo- cation must be made on the basis of geo- graphic probability. Masticophis flagellum fuliginosus (Cope), new combination Baja California Whipsnake Bascanium flafielliforme testaceum: Yarrow, 1883: 112 (part) Bascanion fJagclUforme testaceum: Belding, 1887: 98 Bascanion lateraJe: Cope, 1890: 147 Zamcnis lateralis fulifiinostts Cope. 1895: 679 Bascanion flagellum frcnatum: Van Denl)urgh, 1895: 148 Zamenis flagellum frcnata: Nfocquard, 1899: 323 Bascanion latcralc fuliginosunr. \'an Denliiirgh, 1905: 26; Van Dcnhurgh and Slevin, 1914: 145 Bascanium flagellum: Meek, 1906: 15 (part) Coluber flagellum piceus: \'an Denl)nrgh and Slevin, 1921: 64; Schmidt, 1922: 683: \'an Denburgh, 1922: 664 (part); Klaul)er, 1942: 88 (part) Masticophis piceus: Ortenbnrger, 1928: 125 (part); Lin.sdale, 1932: 375 Masticophis flagellum fremitus: Ortenburger, 1928: 112 Masticophis flagellum frenatum: Linsdale, 1932: 375 Coluber flagellum frenatum: Mosauer, 1936: 16 64 Tiilane Studies in Zoology and Botany Vol. 16 • 4 ■'- -■ . - . — .>-v y^.: Figure 18. Pale phase of Masticophis flagellum fuliginosiis from 7 miles N San Pedro, Baja California del Sur, Mexico. Masticophis flageUum piccus: Brattstrom and Warren, 1953: 179 (part); Cliff, 1954: 77; Murray, 1955: 45; Leviton and Banta, 1964: 150; Stebbins, 1966: 150 (part) Syntypes.—USNM 15135-15136, collected on Santa Margarita Island, Baja California del Sur, Mexico, on 2 May 1888 by the crew of the U. S. Fish Commission vessel Albatross. Diagnosis. — A subspecies of Masticophis flagellum with two color phases, one having a yellow or light gray dorsum with a zig- zag pattern of black bands along the body and wider dark brown bands on the neck (Fig. 18), and the other having a dark grayish brown dorsum with a lineate pat- tern of variable length on the lateral portion of the dorsum, and a variable amount of cream coloration on the venter. Range. — All of the peninsula of Baja California except for the portion east of the San Pedro Martir Mountains and north of Bahia San Felipe. The dark phase extends into the southern portion of San Diego County, California (Figs. 2 and 3). Description of the Syntypes. — Scutellation (the first set of numbers applies to USNM 15135, the second to USNM 15136) : supra- labials 8-8, 8-8, the fourth and fifth entering the orbit in both; infralabials 10-10, 10-10, four touching the anterior chin shields and the fifth the largest in both; loreal 1-1, 1-1; preoculars 2-2, 2-2; postoculars 2-2, 2-2; temporals 8-8, 7-7; scale reduction pattern (of USNM 15135) 15 3 + 4(7) 3 + 4(108) 19 17 3 + 4(8) 3 + 4(112) 14 6 + 7(118) 6 + 7(124) 13; ventrals 192 (Cope gave an incorrect count of 201), 190 (Cope's count is 205); tail incomplete in both. Body lengths are 809 mm and 670 mm, respectively, tail incom- plete in both. Cope (1895) described the color pattern as follows: "Color above blackish-brown an- teriorly, becoming lighter posteriorly to the end of the tail. The dark color extends on each end of the gastrosteges to the angula- tion throughout the length, and in the younger specimen [USNM 15136], fading out beyond the middle of the length. Ground color of belly yellow. In the larger specimen [USNM 15135] the black-brown predomi- nates on the inferior surfaces, yielding gradu- ally to the ground color, which predominates on the inferior surface of the tail. A yellow spot on the preocular; and in the younger specimen on the postoculars and labial plates. Gular and genial plates yellow spotted in the younger specimen, nearly uniform dark brown in the older. On the anterior part of the body of the younger specimen the lateral scales to the third and fourth row have brown shades, with an obscure trace of cross-banding. On the same speci- men near the middle of the body, there are two pale half-cross-bands near together. No. 2 Coachwhip Snake, Masticophis flagclh/m 65 In the same, the center of each parietal plate ternate in position with those above and is brown." below, the placement of this pigment pro- To this description I can add that both duces a zig-zag pattern, specimens show a light flecking throughout The dark phase of juliginosits may be the length of the body and on the side of distinguished from the dark phases of bf)th the neck, this being better developed in piceus and cinguluni by the following char- the smaller of the two specimens. acters: The dorsal coloration is dark grayish Discussion. — Meristic and mensural data brown; the scales of the anterolateral por- for this subspecies based on 92 males and tion of the dorsum have pale lateral edges 70 females may be summarized as follows: that give the impression of a pale, narrow supralabials 7 to 9, infralabials 8 to 11, line along the junction of each scale row preoculars 1 or 2, postoculars 1 to 3, loreal with that above and below; this coloration 1 or 2, ventrals in males 175 to 204, ven- extends for a variable length along the body; trals in females 176 to 205, subcaudals in the head is usually brown, a little paler in males 100 to 129, subcaudals in females coloration than the dorsum; a varying por- 99 to 123, dorsal scale rows 19-17-12 to tion of the venter is cream to yellowish 19-17-13. Body length in males of all ages cream. In contrast, the dark phases of cingu- ranges from 239 to 1320 mm, in females luni and piceus are solid black above and 237 to 1242 mm. Total length in males of the posterior portion of the venter is salmon all ages ranges from 317 to 1729 mm, in pink to salmon red; in these forms the head females 311 to 1676 mm. Tail length/total is the same color as the dorsum, length ratios in males range from 0.209 to The following description of the color 0.288, in females 0.224 to 0.295. pattern of a specimen from Isla Carmen, Because of demonstrable and consistent Gulf of California (JFC 65-204), is char- differences in coloration and pattern, I rec- acteristic of the pale phase of this subspecies, ognize the populations of Masticophis flagel- The greater part of the dorsum is yellowish lufn inhabiting the greater part of the pe- tan. The ground color of the neck, however, ninsula of Baja California as distinct, and, is olive green and the scales are heavily therefore, resurrect the name fuliginosus for speckled with black. There is an orange suf- this subspecies. The race was described by fusion on the first dorsal scale row and to Cope (1895) from the Isla Santa Margarita, some extent on the second. The venter is but he assigned it to the species lateralis, pale yellow anteriorly and lemon yellow It was placed in the synonymy of Coluber posteriorly. There is a double row of ir- flagellum piceus ( = Masticophis flagellum regularly shaped cinammon blotches on the piceus) by Schmidt (1922), who noted the neck. The head is grayish brown dorsally. resemblance of Cope's specimens to snakes The lateral area of the head is similar with from the mainland of Baja California. The white blotches on the loreal, upper pre- name Zamenis lateralis fuliginosus has re- ocular, and supralabials. The chin is white mained in the synonymy of M. f. piceus to with brown blotching, this date. The following description is of a dark This subspecies may be distinguished phase fuliginosus from 21 to 22 miles S La from M. f. piceus and all other races of Paz ( LSUMZ 12630). The dorsum is dark Masticophis flagellutn by the following char- brown grading to brown on the extreme acters: The dorsal coloration of the pale posterior portion of the body and tail. The phase is some shade of yellow ( usually first dorsal scale row is a shade paler. The bright yellow) or light gray (pink to red venter is dark brown on the throat grading in piceus); the dorsal pattern consists of to creamy white posteriorly. The head is a series of transverse zig-zag bands, between dark brown above, brown laterally, with a which the ground color is uniformly pale white blotch on the sixth supralabial. The (in piceus there is an alternating series of chin is brown with cream blotching, which red and cream bands between the dark extends to the posteroventral edge of the bands); and the dark bands in fuliginosus last supralabial. are formed by dark markings at the apex Although there is some degree of vari- and anterolateral edges of each scale. Thus, ability in the intensity of the color pattern since the scales of each dorsal scale row al- at any one stage in a snake's life, there is 66 Tulane Studies in Zoology and Botany Vol. 16 a definite ontogenetic darkening of the pat- tern from juvenile to adult. This change is illustrated by the following description of a series of dark phase jidiginosus from vari- ous localities in Baja California del Norte in the collection of Mr. Joseph F. Copp. New-born specimens have essentially the same color pattern as do the young of "red phase" piceus, but the color is much darker. There are dark brown bands on the neck separated by cream bars. The rest of the dorsum is covered with a mottling of dark and light brown. The venter is cream with dark brown spotting on the neck. The head is brown with cream blotching on the side. The chin is cream with brown mottling. As a snake grows larger (as exemplified by JFC 441 from 2.2 miles S Colnett, Baja California del Norte, snout-vent length 472 mm) the entire dorsum begins to darken. The mottling is less evident and the pale interspaces between the neck bands are re- stricted to the side of the neck. Cream blotching on the side of the head is less extensive. The venter is much the same at this stage as in new-born animals. At a snout-vent length of 967 mm ( JFC 60-94 from 4.2 miles S Rosarito, Baja Cali- fornia del Norte ) , a snake is much darker. The dorsum is very dark brown from scale rows five or six to the middorsal row. Rows one to five or six have a lineate pattern that is more diffuse posteriorly. The cream inter- spaces on the neck are still visible although almost obscured by the lineate pattern. The venter is cream with a double row of light- centered brown spots. The head is darker and the brown blotching on the side of the head and chin is darker and more extensive. As a snout-vent length of 1029 mm (jFC 60-100 from 2 miles N Arrovo San Telmo, Baja California del Norte) the dark colora- tion of the dorsum is more extensive, tend- ing to obscure the lineate pattern on the side of the neck. Spotting on the venter is more prominent and the spots on the neck show indications of fusing together. The brown coloration on the side of the head and chin is more extensive. At a snout-vent length of 1044 mm (JFC .398 from the San Telmo Valley, 3 miles E San Telmo, Baja California del Norte) the cream blotching on the side of the head has almost disappeared and the chin and throat are heavily blotched with grayish brown. At a snout-vent length of 1068 mm (JFC 59-73 from Punta Cabras, Baja California del Norte) the lineate pattern on the side of the neck has virtually vanished, and there is only a slight indication of it on scale row one. At a snout-vent length of 1267 mm (JFC 60-110 from the north base of Punta Banda, Baja California del Norte ) the side of the head and chin have very little cream blotch- ing. In summary, as a juvenile snake of the dark phase of fuliginosus attains adulthood, there is a progressive darkening of the dor- sum, head, throat, and chin, and an intensi- fication of the spotting on the venter. The dorsum of the light phase of fuligi- nosus is either pale to dark yellow, pale tan to tan, or pale gray. Some specimens that have been long preserved are ivory-white in coloration; this may be the result of fading from an original pale gray coloration. In a series of 14 specimens from Mira- flores, 6 are yellow, 2 pale gray to grayish tan, and 6 are ivory-white. The number of specimens of each color type that I have examined from several localities in Baja California are as follows: La Paz — yellow (2), tan (1), pale gray to dark gray (3); Agua Caliente — yellow ( 1 ) , tan ( 2 ) ; San Jose del Cabo — yellow ( 2 ) , tan to grayish tan ( 3 ) ; Cabo San Lucas — yellow ( 3 ) , tan (10), pale gray (2); Santa Anita — yellow ( 1 ) , tan ( 3 ) . That this difference in colora- tion is not due to variations in methods of preservation or storage of specimens is evi- dent from the presence of both yellow and pale gray specimens in series collected at the same time. There is also variability in the amount of dark pigment on the body (Fig. 19). Distribution of the dark and pale phases of the subspecies is shown in Figure 20, in which it is obvious that the black phase is more common in the northern portion of the peninsula. A single specimen of pale- phase fuliginosus (USNM 37551) is avail- able from that portion of the peninsula. It was collected in Trinidad Valley at the northwest base of the San Pedro Martir Mountains by Nelson and Goldman. On the other hand, the pale phase is the more abun- dant form in the southern part of the pe- No. 2 Coachtvhip Snake. Masticophis flcii^ell//?>i 67 A v: .-V •, •» Figure 24. Dorsal view of a juvenile speci- men of Masticophis flagellum from Hacienda Simon, 7.6 miles N Culiacan, Sinaloa, Mexico (UMMZ 120233). jlagelh/m in northern Sinaloa. I had been under the impression that the specimens from this area were all of the pattern D type of M. f. cingulum, i.e., unicolored. How- ever, during the final stages of this study I received several specimens that did not fit my concept of cingulum. This material in- cludes individuals of all size groups. As previously noted, southern popula- tions of cingtihmi lack the banded pattern that is typical farther north, and adults are uniformly tan or brown dorsum ( in pre- served material ) . I have seen no living speci- mens from southern Sonora, but Joseph F. Copp has reported that specimens from the area are nearly uniform red. In Sinaloa the large adults are nearly uniform tan in color ( in preservative ) , but have some small gray or black spots on many of the dorsal scales (LACM 6995-96). Subadult specimens (AMNH 90728-32; KU 67688) usually have more pronounced spot- ting and, in addition, they retain the typical juvenile head pattern of the piceus group until they attain a greater body size than is the case in either cingulum or piceus ( Fig. 23). The dorsal head pattern is much more pronounced than in juvenile cingulum. The border of each dorsal head scale is outlined with white or cream. The area immediately within the cream area is dark brown and the middle portion of the scale is pale brown. The juveniles, although they have the typical piceus group juvenile pattern, have brighter head markings and a more spotted dorsum. The following description is of the head pattern of UMMZ 120233, a juvenile from Hacienda Simon, 7.6 miles N Culiacan, Sina- loa (Fig. 24). The internasals and pre- frontals have wide cream borders anteriorly and medially. The greater part of each scale is dark brown; the posterior edge is tan. The frontal is edged with cream anteriorly and tan laterally. Inside this area there is another narrow region of dark brown. The middle of the frontal is pale brown. The supraoculars are edged with cream an- teriorly and laterally and tan posteriorly. Most of the rest of the scale is dark brown except for the median and outer lateral por- tions, which are light brown. The parietals have a cream spot anteriorly and a continua- tion of the cream stripe on the upper post- ocular onto the anterolateral area. The lat- eral edge of the parietal is edged with dark brown. The ventral portion of the rostral is cream. The upper half is brown with a central light brown blotch. The anterior nasal is mostly brown except for the an- terior edge, which is cream. The posterior nasal is dark brown dorsally and ventrally with a central cream area. The loreal is dark brown with an anteromedial cream blotch. No. 2 Coach whip Snake, Masticophis flagellmn 73 The upper preocular is dark brown along, and dorsal to, the canthus rostraUs; the ventral portion is cream. The lower pre- ocular is cream anteriorly, dark brown pos- teriorly. The upper postocular is dark brown with a central cream stripe. The supralabials are cream with dark brown blotching. The temporals are cream with dark brown blotch- ing- These specimens differ from M. f. cingu- h/m in having, as adults, a patternless dor- sum, except for some small spots on the dorsal scales, and in having the juvenile head pattern accentuated and retained in larger individuals. Subadult specimens from Sinaloa are patternless, whereas adult picei/s are banded. In addition, juveniles from Sina- loa are much more brightly marked than juvenile piceus and the dorsum is more spotted. These specimens from Sinaloa may repre- sent a distinct pattern form that deserves subspecific recognition. I have seen insuf- ficient material, however, to make such a decision at this time. Key to the Subspecies of Adult Masticophis flagelh/f/i This key is designed so that all pattern phases of the subspecies of Masticophis jIageUum may be identified. 1. Each dorsal .scale (at least on the an- terior portion of the body) with a dark brown to black medial line; mid- portion of venter bright yellow; sub- caudals and posterior portion of venter with a salmon pink to red posterior border which does not fade in pre- servative Masticophis flagellum lincatultis Pattern not as above 2 2. Anterior 1/3 to 1/2 of dorsum dark to very dark brown (some specimens nearly black), posterior 2/3 to 1/2 brown to tan ( in black specimens, the posterior portion of the venter and underside of tail are cream as opposed to the pink coloration (in life) of this area in black M. f. piceus and M. f. cingulum ) Masticophis flagellum flagellum Color pattern not as above 3 3. Some portion of dorsum red or pink in life (tan or li^ht brown in preser- vative) 4 Dorsum not red or pink 10 4. Anterior portion of dorsum solid black, posterior portion red 5 Pattern not as above 6 5. Posterior 1/3 to 1/2 of body with a pattern of black bands separated In- light red (tan in preservative) inter- spaces; sometimes a narrower, black band bisects the light interspaces ... Masticophis flagellum cinguluui (pattern F) Posterior 1/3 to 1/2 of l)ody with a variegated pattern of red and l)lack scales with no division into dark bands and narrow, light interspaces Masticophis flagellum piceus ( intermediate phase ) 6. Dorsum dark red to dark redtlisli- brown, this coloration being broken into long hands (11 to 14 scales long ) by light (pink in life) interspaces 5 to 6 scales in length. A dark narrow liand usually bisects each light interspace. This type of pattern usually extends the length of the body (pattern class A ) l)ut some specimens have this band- ing reduced and confined to a variable amount of the anterior portion of tlie body (pattern class B) 7 Dorsal banding, if present, confined to neck 8 7. Some black scales on neck — Masticojihis flagellum cingulum ( pattern type E ) No black scales on neck Masticophis flagellum cingulum ( pattern classes A and B ) 8. A single light hand across nape __ Masticophis flagellum cingulum (pattern C) Pattern not as above 9 9. Dorsum with relatively long (3 to 4 scales long) black, gray, or brown ( also sometimes the same color as the dorsum, outlined anteriorly and pos- teriorly with darked pigment ) bands on neck and a liglit, dark-bordered stripe through loreal - - Masticophis flagellum piceus (red phase) Dorsum uniformly pink or salmon pink with slight banding on neck (bands, when present, narrow, about 1 scale long); no light, dark-bordered stripe through loreal - __.. Masticophis flageUum testaceus (pink phase) 10. Dorsum tan to light brown, yellow or light gray, or light yellow to light olive yellow H Dorsum dark gra> ish-brown to black 17 11. Dorsum tan to light I)r()vvn 12 Dorsum yellow, light gray, or light yellow to light olive yellow 16 12. Remnants of juvenile pattern present: dark juvenile head markings of flagel- lum group present on dorsal head scales; bands on neck dark brown and well defined Masticophis flagellum flagellum ( pale phase ) Dorsal pattern not as al)ove 13 13. Dorsum umforni tan to light brown (at least in preser\ati\e) with no pattern 74 T/dane Studies in Zoology and Botany Vol. 16 Masticophis flageUum testaceus (patternless phase), Masticophis flagel- lum cingulum (patternless phase), and Sinaloan specimens Dorsum tan with some type of pat- tern 14 14. Dorsum uniform tan to light brown with a narrow, light band across nape of neck ... Masticophis flageUum cinguUim (pattern C) Dorsal pattern not as above 15 15. Dorsum with narrow ( usually 1 and sometimes 2 scales long ) brown to dark brown bands on neck and forebody; dorsum of head more or less unifonn in coloration __Masticoph is flageUum testaceus ( narrow-banded phase ) Dorsum tan with wide, brown bands separated by narrower, lighter inter- spaces; wide bands sometimes com- posed of a series of dark brown narrow bands which are separated by a much lighter, shorter series of narrow bands Masticophis flageUum testaceus ( wide-banded phase ) 16. Dorsum yellow or light gray with nar- row, zig-zag black bands on dorsum, and with wider dark brown bands on neck; a double row of brown to dark brown spots on anterior portion of ven- ter Masticophis flageUum ftdiginosus (pale phase) Dorsum light yellow to light olive yel- low; pattern absent or with obsolete bands on neck more or less the same color as the ground color; no spotting on anterior portion of venter Masticophis flageUum ruddocki 17. Dorsum and head completely black; posterior portion of venter and under- side of tail salmon pink to red Masticophis flageUum piceus (black phase) and Masticophis flageUum cingulum (black phase) Dorsum dark grayish-brown with indi- cation of light lines on a variable amount of the lateral area of the dorsal scales; head usually somewhat lighter than rest of dorsum; venter usually completely cream with a double row of dark spots on anterior portion ( specimens with a variable amount of dark pigment on anterior portion of venter still have cream coloration on posterior portion of the body) Masticophis flageUum fuliginosus (dark phase) Significance and Evolution of Color Pattern Masticophis jlagelhnn is one of the most variable snakes in North America. Not only is the species divisible into geographic races, but almost all the subspecies exhibit pattern variations. Only two, AI. /. lineatulus and M. f. ruddocki, do not. Also, most of the subspecies exhibit an ontogenetic change in color pattern. Only AI. /. testaceus, of the testaceus group, and the red phase of Al. /. piceus and the pale phase of AI. /. fuligino- sus, both of the piceus group, show very little change in color pattern from juvenile to adult. The types of variation in adult pattern within the subspecies of Masticophis flagel- lum fall into four categories. These are: ( 1 ) clinal variation, ( 2 ) random variation, ( 3 ) non-clinal geographic variation, and (4) polymorphic variation. Clinal variation in color pattern is exhibited by AI. /. flageUutn and AI. /. cingulum. In Al /. flageUum the color pattern is paler (has less dark brown pigment on the anterior portion of the body) in the southeastern portion of its range and is darker in the northwestern portion of its range. In AI. /. cingulum there are two, more or less parallel clines within the range. First, there is a clinal trend toward loss of the banded pattern ( pattern A ) , found most commonly in the northern and northeastern portions of the range, to the unicolor pattern (pattern D), found most commonly in the southern por- tion of the range. Second, there is another clinal trend toward progressive melaniza- tion of the banded pattern (pattern A) of the northern and northeastern portion of the range to produce the black pattern (pat- tern G), found in the central and eastern portions of the range. This deposition of black pigment takes place from anterior to posterior, and some specimens are totally black dorsally, thus paralleling the type of pat- tern seen in the black phase of AI. /. piceus. Random variation is seen in all subspecies, but it is most conspicuous in M. f. testaceus, in which the variation in color pattern is discontinuous, i.e., there are three dis- tinct patterns found in varying proportions throughout the range. Geographic variation within subspecies limits is seen in M. f. jlagelhnn and AI. /. testaceus. In AI. /. flageUum there is a pale tan phase centered in Florida, the adult pat- tern of which closely resembles the pattern of all juveniles of the testaceus group gen- erally, and the pattern of juvenile flageUum particularly. This phase has a clearly de- fined, albeit discontinuous, range in portions No. 2 Coachwhip Snake, Masticophis flagdlmn 75 of Florida and Georgia ( a single comparable stated "the genes involved in polymorphism specimen from North Carolina was also have, in general, conspicuous discontinuous seen). In M. f. testacei/s there is a pink effects, and different genotypes (except phase, which differs from the more com- some heterozygotes ) can be distinguished mon and widespread tan form. This phase phenotypically." If the intermediate is con- inhabits three disjunct areas in Trans-Pecos sidered a heterozygote, resulting from the Texas, eastern New Mexico, and eastern crossing of a homozygous pale phase snake Colorado. with a homozygous black phase snake, then Polymorphism is seen in M. f. piceus and the situation cannot be designated as bal- M. f. juliginosus. Both subspecies have a anced polymorphism, which is maintained pale (red in piceus, yellow, tan, or light by "overdominance" of the heterozygote gray m full ginosus) and a dark phase (black (Mayr, 1963, p. 221). In fact, the inter- in piceus, dark grayish-brown in fuligino- mediates appear with a much lower fre- sus ) . Intermediate specimens are more com- quency than either the pure dark or pale mon in the case of piceus, less so in fuligino- phases, at least in the peninsula of Baja sus. There is a dearth of information on California. Thus the usual solution to the the basis for this polymorphism. It is not problem of the maintenance of poly- known whether dark phase adults always morphism does not seem to apply here, give birth to dark phase young and light This situation is similar to that described phase adults to light phase young, or for the screech owl (Oius asio) by Owen whether both might appear in a single (1963). Two phases of the screech owl, a brood of either phase as is the case in Lam- gray one and a rufous one, exist over the propeltis getulus calif orniae ( Klauber, portion of eastern United States lying east 1944). There is no information, other than of meridian 104° W. Intermediate specimens that which I have presented in discussing also occur, as is the case among the whip color pattern variation, on the relative fre- snakes. The relative frequency of rufous quency of each color pattern in any given screech owls varies geographically, as does population. It is known that the range of the the relative frequency of intermediate birds, black phase of AI. /. piceus is centered Owen suggested that this type of poly- around Tucson, and that radiating from morphism, which he preferred to call bi- that central point the phase becomes less modal variation, is the result of a balance frequent in occurrence. It is also known of selective forces acting on each phase and that within its range the black phase is most that the presence and maintenance of these common, the intermediate phase next, and two phases "represent balanced adaptations the pale phase least common. of the forms to varying environmental con- Within the subspecies fuUginosus the ditions." He goes on to say that "the exis- picture is less clear, largely because of in- tence of two distinct forms with few inter- sufficient material. Are there areas where mediates throughout much of the range of only the pale phase occurs, as is suggested the screech owl in eastern North America by the specimens available from the ex- is indicative of selection for bimodal varia- treme southern cape region of the peninsula tion. In most areas, selection must be con- of Baja California and some of the gulf tinually operating against intermediates, but islands? Are there areas where only the in Florida, where intermediates are frequent dark phase occurs, as is suggested by ma- . . . , there is presumably a relaxation of terial from the northwest coast of Baja Cali- selection pressure. The result of this is that fornia del Norte and others of the gulf while all six color categories [a pattern islands? If so, why is this the case and why classification used by Owen to determine is it that both phases occur in some areas? the frequencies of intermediates] occur in What is the genetic basis of this pattern Florida, the population is extremely vari- dimorphism? It would appear that neither able, but not polymorphic. Hence in the in the case of piceus nor that of fuliginosus screech owl the unusual situation exists in is it based on a single pair of alleles, for which polymorphism for color and pattern intermediate specimens do exist, which sug- is maintained over a large geographical area gest the action of modifier genes in produc- (eastern North America), while in a much ing the adult pattern. Mayr (1963, p. 151) smaller area (Florida) the complete range 76 Tidaiie St //dies in Zoology and Botany Vol. 16 of phenotypes occur, but the population is a pale blotch on the upper preocular. In not polymorphic." West of the 104th contrast, in the eastern group, the pale meridian all the birds are gray. spots on the nasal and loreal scales are not Although the characteristics of the poly- in contact but are separated by a thin stripe morphism exhibited by the subspecies pice/ts of brown pigment along the posterior edge and f/diginosm are less well documented, of the posterior nasal. Also, the pale spots there appear to be some differences from on the nasal and loreal are not oriented the review presented by Owen for the longitudinally, and the spot on the loreal screech owl. Some of these are due to the is not in contact with the spot on the upper smaller range inhabited by the black phase preocular. There is considerable irregular whipsnakes and to the smaller number of cream blotching on the temporals of the specimens available for study. In f//liginos//s, western group, which is less evident or ab- however, it appears that intermediates are sent in the eastern group, few in number, thus presenting a condition In both groups there is a subspecies that approaching bimodal variation. In pice/ts, retains the juvenile pattern essentially un- on the other hand, intermediate specimens changed to the adult stage. In the eastern are of rather frequent occurrence and there {testace/is) group it is the subspecies tes- would seem to be little selective pressures tace//s, and in the western (piceus) group acting against the maintenance of this type it is the subspecies pice/ts. The other sub- of pattern. Also, in the area where the black species of each group have departed in vary- phase of pice/is occurs, it is the most common ing degrees from this pattern, at least in phase, and the light phase is the least com- the adult color pattern. mon. Thus, at least in the case of piceus. Thus, the division of the species flagelh/m there would appear to be little selection for into two stocks appears to have taken place bimodal variation. at an early time, inasmuch as the basic pat- In any event, to place a name on the terns of the piceus and testaceus groups are type of variation exhibited by piceus and markedly different. None of the juvenile fuliginosus would be no solution to the patterns of the subspecies in either group problem of why, in certain areas, a poly- shows any approach in basic pattern to ju- morphic phase is maintained. If there are veniles of the subspecies of the other subtle differences in niche preference, it is group. not presently known what they might be. Within the testaceus group, the sub- There is, in fact, very little information on species testaceus is the central form from any aspect of the ecology of Masticopbis which the other two subspecies, flagel- flagell//m. I um and lineatulus, have diverged. M. It is evident from a study of juvenile pat- /. flagellum has invaded the eastern tern, that there are two main evolutionary United States and become adapted to lines within the species jlagelh/m. One, an the Austroriparian type of vegetation. M. eastern group of three subspecies {jlagellum, f. lineat/this has invaded the Chihuahuan testaceus, and lineatulus) , is characterized Desert of the northern portion of the Mexi- by having a juvenile pattern composed of a can Plateau. These two subspecies have di- tan dorsum with narrow, dark crossbands verged from more or less opposite ends of ( usually 1 scale long, sometimes 2 ) sepa- the range of the testaceus-Wkc ancestor and rated by wider interspaces (Fig. 25). The have little in common with each other, other, a western group of four subspecies flagellum having developed a dark brown {piceus. cingul/im, r/iddocki, and j/iligino- anterior body color and lineat/ilus a lineate s/is), differs from the eastern group in hav- dorsum and a salmon-red venter, ing relatively long neck bands (3 to 4 Within the pice/is group the subspecies scales in length ) and narrow ( 1 scale in piceus, or its ancestor, is the central stock length) interspaces (Fig. 25). The mem- from which the other subspecies have di- bers of this western group also have verged. One subspecies has invaded Mexico a pale streak through the nasal and loreal west of the Sierra Madre Occidental and scales. This streak usually is straight- evolved a different type of banded pattern edged and longitudinal, and usually extends from that seen in piceus. This is the sub- completely through the loreal, joining with species cing/il/im. The subspecies ruddocki I No. 2 Coach whip Snake, Mastic op his jlagelhm 11 ABC Figure 25. Dorsal view of the head of juvenile Masticophis flagelliirn flagellum (A), M. f. tes- tacctis (B), and M. /. piceus (C). has invaded the central valley of California and developed a pattern lacking the type of neck banding seen in piceiis and having a different ground color. The subspecies \nliginosus has invaded the peninsula of Baja California and developed a basically different ground color and a slightly dif- ferent type of banding on the dorsum. Variation in Scutellation AND Proportions Head Scutellation Siipralahials. — The number of supralabials is normally 8, the fourth and fifth entering the orbit. Variation from this number is infrequent. Only 7.9% of the specimens examined has a count other than 8-8. De- partures from the usual 8-8 condition in- volve the fusion or division of scales. In- crease in the number of supralabials most often involves the division of the third supralabial (82.1%), but may also involve the division of the fourth ( 10.7% ), second ( 3.6% ) , eighth ( 2.4% ) , or fifth ( 1 .2% ) . Decrease in the number of supralabials most often involves the fusion of the second and third supralabials (52.0%), but may also involve the fusion of the seventh and eighth (36.0%), or the fourth and fifth ( 12.0%). One specimen (TNHC 29117) has the second and third supralabials fused and the seventh divided giving the normal count of eight. The third and seventh supralabials are divided in JFC 63-150. In TNHC 7341 only the fifth supralabial enters the orbit. What apparently has taken place is that one of the anteocular supralabials is divided (probably the third) and the fourth and fifth are fused. In UCM 31533 the fourth supralabial does not enter the eye on the left side and is not present on the right. NTSU 193 has the seventh supralabial fused with one of the temporals. Variation in these scales is shown in Tables 4 and 5. Infralabials. — The number of infralabials is most often 10 on each side of the head 78 Tuhwe Studies in Zoology and Botany Table 4. Variation in number of supralahials in Masticophis flagellum. Vol. 16 Subspecies 7-7 7-8 8-8 8-9 9-9 Total flagellum 3 321 16 2 342 tcsfacctis 4 10 499 27 5 545 lineatulus — 29 4 1 34 piceiis — 1 238 12 — 251 cingtthnn — 1 74 2 1 78 niddocki — 9 3 — 12 fidigiiwsus 2 2 49 9 — 62 Total 6 17 1219 73 9 1324 Table 5. Variation in number of supralabials entering orbit in Masticophis flagellum. Subspecies 3+4 3+4 3+4 4+5 4+5 4+5 4+5 5+6 5+6 5+6 4 5 5 4 4 Total 4+5 flagellum ^ 3 321 16 2 _ 342 tcstaceus 1 5 491 27 3 2 2 1 532 lineattdus — — 29 4 1 — — — 34 })iccus — 1 238 11 — — — — 250 cingulum _ _ 75 4 1 80 ruddocki _ — 9 3 1 _ _ _ 13 fuliginosus 1 3 50 9 — — — — 63 Total 2 11 1213 74 8 2 2 1 1313 Table 6. Variation in number of infralabials in Masticophis flagellum. 8- 9- 9- 9- 10- lo- 10- 11- 11- 12- 11- 8- 8- Subspecies 10 9 10 11 10 ll 12 11 12 12 13 9 11 Total flagellum 1 2 22 1 159 80 8 35 8 1 1 318 tcstaceus 1 17 54 8 248 121 3 29 6 1 1 1 490 lineatulus — _ 6 1 14 5 1 — _ — — _ — 27 ptceus — 1 12 4 101 74 5 39 5 _ _ — _ 241 cingulum — — 4 _ 31 20 3 11 2 _ _ _ — 71 ruddocki — — 1 _ 3 3 1 2 — — — _ — 10 fuliginosus — 6 8 — 36 9 _ 2 _ _ _ 1 _ 62 Total 2 26 107 14 592 312 21 118 21 2 1 2 1 1219 Table 7. Variation in number of infralabials contacting anterior chin shields in Masticophis flagellum. Subspecies 3-3 3-4 4-4 4-5 5-5 Total flagellum 1 283 26 4 314 testaceus — 4 460 17 1 482 lineatulus — _ 23 3 1 27 piceus 1 — 230 8 2 241 cingulum — 1 68 3 73 ruddocki _ 6 6 12 fuliginosus 7 2 53 72 Total 9 7 1123 63 8 1210 No. 2 Coachichip Snake, Masticophis jlagellum 79 Table 8. Variation in number of loreals in Masticophis jlagellum. Subspecies 0-1 1-1 1-2 2-2 2-3 Total jlagellum 3 341 3 5 1 353 testaceus — 515 15 4 — 534 lincatulus 1 32 — — 1 34 piceus _ 253 1 — — 254 cingulum 1 78 — — — 79 ruddocki 1 11 — — — 12 juliginosus — 69 1 — — 70 Total 6 1399 20 9 2 1436 (48.6%), 4 infralabials being in contact with the anterior chin shields, the fifth the largest. Deviations from this number, how- ever, are manifold ( Tables 6 and 7 ) . The first to the fifth infralabials are relatively stable and most variation takes place be- hind this area. The variation in number of infralabials posterior to the fifth is more erratic and involves the addition, deletion, fusion, and division of small scales. Thus, the pattern is not clear. Division and fusion, however, do take place within this relatively stable anterior zone. Increase in the num- ber of infralabials in this area most com- monly involves the division of the third infralabial (82.4%), but also may involve the division of the fourth (11.8%) or sec- ond ( 5.8% ) infralabials. Decrease in the number of infralabials in this area most often involves the fusion of the third and fourth infralabials (71.4%), but also may involve the fusion of the second and third infralabials ( 28.6% ) . Variation in these scales is shown in Tables 6 and 7. Nasal. — The nasal scale is always divided vertically. A few specimens, however, have the postnasal section divided horizontally. Loreal. — The loreal usually is single on both sides of the head (97.4%). Increase in the number of loreals involves the hori- zontal division of the existing loreal into two parts on one or both sides of the head and the addition of one or more smaller scales, usually between the loreal and the supralabials. Decrease in the number of loreals involves the fusion of this scale either with the prefrontal ( 50.0% ) or with the posterior nasal ( 50.0% ) . Variation in this scale is shown in Table 8. Preoculars. — The number of preoculars usually is 2 (98.9% ). Increase in the num- ber of preoculars involves either the forma- tion of a suture at the junction of the canthal ridge on this scale and the flat vertical por- tion (28.6%), the formation of a suture farther down on the vertical portion of this scale (28.6%), or the splitting off of a scale from the fourth supralabial to form a third preocular-like scale (42.8%). De- crease in the number of preoculars is an uncommon event, but when it occurs it in- volves the fusion of the lower preocular with the third infralabial. Variation in these scales is shown in Table 9. Postoculars. — The numbers of postoculars predominantly is 2 ( 98.0% ) . Increase in the number of postoculars involves the for- mation of a suture either in the middle of the lower postocular or in the middle of the upper postocular, or involves the fusion Table 9. Variation in numbers oj preoculars in Masticophis jlagellum. Subspecies 1-1 1-2 2-2 2-3 3-3 3-4 Total jlagellum _ 342 1 343 testaceus _ _ 528 4 1 — 533 lincatulus 33 — — — 33 piceus 246 3 2 — 251 cingulum 1 _ 77 1 - — 79 ruddocki — — 11 - - 1 12 juliginosus _ 1 70 — — — 71 Total 1 1 1307 9 3 1 1322 80 Tnlane Studies in Zoology and Botany Vol. 16 Table 10. Variation in number of postoculars in Masticophis flagellum. Subspecies 1-1 1-2 2-2 2-3 3-3 Total flagellum 1 1 337 2 1 342 testaceus 5 5 516 3 1 530 lineatiiliis _ — 33 — — 33 piceiis — — 255 1 — 256 cingidutu _ _ 79 — — 79 niddocki _ — 12 — _ 12 fitliginosus 1 — 69 2 4 76 Total 7 6 1301 8 6 1328 of the two normal postoculars and the split- ting off of two scales from the fifth supra- labial. Decrease in the number of postoculars involves either the fusion of the upper postocular with the parietal, the fusion of the two postoculars into one scale, or the fusion of the lower postocular with the fifth supralabial. Variation in these scales is shown in Table 10. Frontal. — The frontal normally is bell shaped. The only anomaly observed in this scale is the presence, in two specimens, of a suture through the middle of the scale dividing it into anterior and posterior por- tions. Internasals. — These scales normally are paired. One specimen (NCSM 5886), how- ever, has the right internasal split into right and left halves and the left internasal fused with the anterior nasal. Prefrontals. — These scales are normally paired. One specimen, however, has the pre- frontal area divided into three scales. Body Scutellation Ventrals. — The number of ventrals ranges from 173 {in M. f. testaceus) to 212 (in M. f. flagellum ) . Ranges for the various subspecies are listed in the appropriate sub- species account. The means of the popula- tions east of the Mississippi River are rela- tively constant. All means fall within the range of 201.0 to 203.7 for males and 196.0 to 201.8 for females. The highest means are in Florida, southern Georgia, Alabama, Mississippi, and western South Carolina (Fig. 26). West of the Mississippi River the mean number of ventrals drops sharply and continues to decrease more gradually through central and southern Texas ( Fig. 27). In western Texas the means again be- gin to rise, abruptly so in eastern New Mex- ico. The mean continues to rise through western New Mexico but again begins to decrease in Arizona, Nevada, Utah, and por- tions of southern California. Both to the north and the south of Los Angeles, San Bernardino, and Riverside counties, Cali- fornia, the mean number of ventrals again increases. Immediately to the south of San Diego and Imperial counties, California, the ventral number decreases, but farther down the peninsula of Baja California the mean number increases, until in the cape region the mean number of ventrals approaches some means for southeastern United States, the area which has the highest mean num- ber of ventrals (Fig. 28). In Sonora the mean number of ventrals increases from north to south, but this cline reverses it- self in northern Sinaloa. To the east of the Sierra Madre Occidental, in Chihuahua, the mean number of ventrals is much lower than to the west and most closely resembles means for central Texas. To the south, in Durango and Zacatecas, the mean is similar to that in Chihuahua but is slightly less than the mean of populations to the east in San Luis Potosi and Tamaulipas. There is a minor amount of sexual dimorphism in ventral number but females often have slightly fewer than males. Anal Plate. — The anal plate is normally divided in this species and only two of the specimens examined have the two halves fused. Subcaudals. — The variational trends in subcaudals are less clear than for the ventrals primarily because only about 48% of the specimens examined have complete tails. The number of subcaudals ranges from 91 (in M. f. flagellum) to 129 (in M. f. fuli- ginosus) . No. 2 Coach whip Snake, Masticophis flagellum 81 Figure 26. Geographic variation of ventral number in Masticophis flagellum in the eastern United States. Black Ijlotches represent the areas from which samples were taken. The upper numbers associated with each blotch are the sample mean and sample size (separated by a dash) for males; the lower numbers present the same data for females. When a line appears in place of either the upper or lower figures, it signifies that no specimens of that sex were present in the sample. Variational trends are similar to those observed for ventrals but are not identical. The mean number of subcaudals east of the Mississippi River is relatively high, but means for various populations are not so constant as are those of the ventrals in this area (Fig. 29). West of the Mississippi River the mean number of subcaudals de- creases rather sharply and continues to de- crease through western Oklahoma and eastern and central Texas ( Fig. 30 ) . Except for an in- crease in the male mean in eastern New Mexico, the decrease continues in western New Mexico and Arizona. The mean in- creases again in southern California, except for populations from the central valley and adjacent areas where there are fewer sub- caudals. The mean continues to increase, with some variation, southward through the peninsula of Baja California; means for the southern portion of the peninsula are the highest anywhere within the range (Fig. 31). In Sonora the mean tends to decrease from north to south, rather than increase as does the mean for the ventrals. This trend continues into northern Sinaloa where the mean is the lowest anywhere within the range, with the exception of Chihuahua 82 Ti/lane Studies in Zoology and Botany Vol. 16 192.3-16- " i^v^^H 191.5-14 r^^^^^ "^194.6-23 193.6-10 Figure 27. Geographic variation of ventral number in Masticophis flagelhim in tlie western United States. Explanation as for Figure 26. where the female mean is lower than that for females from northern Sinaloa. Al- though based on relatively few counts, the mean number of subcaudals for central and eastern Mexico is similar to that for central and southern Texas. Sexual dimorphism in subcaudals is evi- dent, with males having, usually, a higher mean number. The amount of difference between the sexes varies from population to population and ranges from about 2 to -I n < CO O cc S O c 1 i> CO m lajor 1 re- mid- tion. two » 1 1 1-' 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 -*■ 1 1 1 1 ^' 1 1 1 1 1 e n tera nd : due 'ith J= 03 O CD :f +J _; y >H "^ CO CO o i-H o o ^H o in 1—1 ithin cond B-se orsal lorsal ^ 1 1 1 1 -t CO 1 1 d 1 1 -J 1 id id 1 1 1 -; 1 1 1 1 d 1 -r 1 1 1 1 -h' . — 1 ^H ^H p-H ^ CM ^ ^ 3 .2 -3 3 3 •— 4-< H " O CJ CO o Tfco t- in ■■a 1 1 1 1 -t 1 1 I— ( 1 I 1 1 1 1 1 in 1 1 1 1 1 t'ln 1 1 1 1 1 1 CO 1 1^ 1 1 r to subdiv tution appe niddorsal re on, B— seco 8— Fusion O O t-; C0_ CO CO ^H OCMO o r^ in 05 in t-^ 00 ^ 1 1 1 1 in 1 -H 1 1 inooin 1 1 in 1 1 1 1 1 1 1 1 CO 1 1 1 1 ^. •-;-HTt_i>.co e lett on, B on; A orsal ebral "* oiin 00 CO ^ r-^ I 1 d CM 1 t-^CO 1 ic oi d d 1 d in 00 c>i d d in t-^ d -H ^" d t-' CO "* ^ CO CO t^ CO CMTf CM ^ CO CM-* CM in in CD a> OMn CO co^cmco co ula. Thi reducti( reducti st midd paravert p q -H t-; CD q O ^ CM CO O CDOO ^ 00 1; o q -f OI in i> CM r- 00 3 rt «;:: .y. ffl a> lo" c^" CO 00 ic* 1 1 in -J 1 oico' 1 in CO in d 1 i—Hin^d iininocxdoot- 't-^ o S a! 1 = ^^ CM (M ^^ CM CM CO CM CM^ CO F-H p-H CO 1— I^HCM-H 1 — ^1 — ^-^^H(M" — i CM Mi i! -ti J- o ""■ — 3 S *- -0 ''^ '^ OO ■•;=! 12 S 3 *S U •- O O 3 3^ ox; 3 =00 COO CDOOO o in CO O -HO coco O O t--; ^ 00 q ^ CM -? 3 ^ ii-3 <: '^ in d ic 00 00 d 1 d>d> 1 1 CO 1 in d d 1 1 1 CO in 1 d 1 d t-^ d -t t-^ d 1 CM le re ars i r to : redi 7-S ^H CO CM CM ^H O CM^ (-M CM CO ^H 1— I 1—1 CO CM r-^ rt OJ O,-^ . o a-r CS 0) re- tire s n ap ante: Idors areas itage! O I- 00 CD mo 1-H o ot^comcocD q m 1 d 1 d 1 1 1 1 1 in 1 in 1 1 1 1^' in 1 1 1 -J 1 1 d 1 ini>coo6dco 1 iri ^H ^H ^^ -H OI 1— I 1— 1 1— H t— 1 CM 35 _C 3 O ai "^TS O V u t« ri oj Qj t« « CO q -H l-; 00 O CO t-co ot-o CMi-HOOOOCM'1d> o -H o p i> q tp 00 ^_ i> q iin 1^ id idt^cot^codt~^in O .ti 3 ■« iH ^ ., XI .b o o 1—1 1—1 CM "—1 '-H f-H CM CM CM ^ -c.= rt 1 fe c < ^. J5 :^^ ^ H-J.^ ;nZc>5UUQNi/ t ■ tion of the posterior portion ot the body Within the Sonoran Desert, M. . cingu- ^^^ geographically variable, specimens from lum is found in all subdivisions. It occurs ^^^ so,,theastern portions of the range being in the Lower Colorado Valley, the Arizona ^^j^^ ^^^^ specimens from the northwestern Upland, the Plains of Sonora, an area of p^^fions of the range. transition between the very arid type of Masticophis flagellum testaceus has two desert vegetation to the north and the more ^yp^^ of ground color and three types of humid subtropical thorn forest vegetation ^(^Jqj. pattern. One ground color is pink or typical of west coast Mexico, the Central j-gj^ this phase characterizing three disjunct Gulf Coast, an area very similar in vegeta- populations in Trans-Pecos Texas, eastern tional characteristics to that of the eastern New Mexico, and eastern Colorado. A tan coast of Baja California below Bahia de Los ground color is found throughout the re- Angeles, and the Foothills of Sonora, a fairly mainder of the range of the subspecies. The rugged area with elements of both the des- three types of color pattern ( a unicolor dor- 94 Tidane Studies in Zoology and Botany Vol. 16 sum, a narrow-banded dorsum, and a wide- banded dorsum) occur throughout the range of Al. /. testaceus. Masticophis flagelhnn Uneatidiis is one of two subspecies that shows little pattern variability. Intergradation between M. f. lineatulus and M. f. testaceus, and between M. f. Uneatuh/s. M. f. piceus. and M. f. cingu- luni takes place over a large area in western New Mexico and southeastern Arizona, re- spectively. Masticophis jlageUinn piceus has two phases, one black, the other red. The black phase occurs in south-central Arizona and is most common within a 20-mile radius of Tucson, Pima County. Masticophis jlagelhnn ruddocki is the other subspecies that shows little pattern variability. Intergradation between M. f. ruddocki and M. f. piceus occurs in low montane passes at the southern end of the San Joaquin Valley. Masticophis flagellum cingulum, as en- visioned in this study, has a range that en- compasses all of Sonora, with the exception of the Gran Desierto of the northwestern portion of the state, and extreme south- central Arizona. This subspecies exhibits considerable variation in color pattern, rang- ing from a banded phase to one in which the dorsum is uniformly black or red. These pattern changes are clinal in nature. The subspecies name Masticophis flagel- lum fuliginosus, a new combination, is ap- plied to the form inhabiting the peninsula of Baja California. This subspecies has a pale and a dark phase, each of which ex- hibits variability in both color and pattern. The two phases occur on the islands on both sides of the peninsula. A distinctive, but poorly known, pattern variant occurs in northern Sinaloa. Ventrals and subcaudals exhibit both geo- graphic and individual variation. The sub- caudals also are subject to sexual dimorphism to a slight degree. The scale reduction pat- tern exhibits considerable individual varia- tion, but certain aspects appear to have geo- graphic significance. Only individual variation is exhibited by the head scutella- tion. Acknowledgments During the course of this study I have been aided by a number of people who have lent me specimens or who have provided me with in- formation or ad\'ice. First of all, I would like to thank my former major professor. Dr. Doug- las A. Rossman, who has always encouraged my research on this and other topics and given freely of his advice. I should like also to thank the other memliers of my committee, Dr. George H. Lowery, Dr. J. Harvey Roberts, Dr. Albert H. Meier, and Dr. Walter J. Harman. To the following people I would like to extend my sincere gratitude for their patience with my requests for the loan of large nimibers of a very large snake and/or information or advice: Steven C. Anderson, Walter Auffen- iaerg, Richard J. Baldauf, Charles M. Bogert, W. Glen Bradkn, Br\ce G. Brown, E. Milb>- Burton. Francis M. Bush, Janalee P. Caldwell, William J. Cliburn, Joseph F. Copp, Peter Diekiewicz, William G. Degenhardt, James R. Dixon, Charles L. Douglas, Neill H. Douglas, William E. Duellnian, Harold A. Dundee, B. Gandy, Glenn Gentry, Bryan P. Glass. Harvey L. Gunderson, Donald E. Hahn, M. Max Hens- ley, M. B. Keasey, HI, Edmund D. Keiser, C. A. Ketchersid, John P. Kerr, F. Wayne King, John M. Legler. Alan E. Leviton, Ernest A. Liner, Charles H. Lowe, Jr., Edmond V. Malnate, T. Paul Maslin, Clarence J. McCoy, John S. Mecham, John D. Miller, Robert H. jvlount, Michael \l. Ovchynnyk, William M. Palmer. James A. Peters, Neil D. Richmond, Douglas A. Rossman, Norton M. Rubenstein, Allan J. Sloan, Hobart M. Smith, Henry M. Stephenson, Wilmer W. Tanner, Ray C. Tel- fair, HI, Robert A. Thomas, Donald W. Tinkle, Charles F. Walker, Ernest E. Williams. Ken- neth L. Williams, and John W. Wright. I would also like to thank the following in- dix'iduals for contributing information useful in this study: Andrew Arata, Ralph W. Axtell, Joseph R. Bailev, C. L. Baker, Rollin H. Baker, Roger W. Barbour, Albert P. Blair, H. T. Boschung, Jr., James L. Christiansen, Joseph T. Collins, James Dobie, Arthur C. Echternacht, Richard Etheridge, Denzel E. Ferguson, T. P. Haines, Laurence M. Hardv, Robert F. Inger, James C. List, Wilfred T. Neill. Brent B. Nickol, Kenneth S. Norris, Gerald G. Raun, R. J. Robel, James T. Tanner, and Richard Thomas. I want sincerel>- to thank Howard W. Camp- bell, Joseph F. Copp, and my former fellow graduate students, Richard M. Blaney, Donald E. Hahn. Edmund D. Keiser, R. Earl Olson, Gerald C. Schaefer, and Kenneth L. Williams, for gi\'ing freely of their knowledge concerning whipsnakes. I would also like to thank Mr. and Mrs. Roger Conant for the use of two photographs from their joint photographic collection and Richard M. Blaney, William G. Degenhardt, Howard K. Gloyd, Gerald C. Schaefer, and Robert G. Webb for the use of photographs from their respective collections. I am grateful to the late Laurence M. Klau- ber for the loan of a large amount of data compiled Iw him during his study of western Masticophis flagclUtin. No. 2 Coachwhip Snake, Masticophis flagellum 95 My sincere thanks go to Anthon\ \V. Ro- National Park, Kentucky. Herpetologica, 5: mano and Robert M. Stabler for the gift of 21-22. specimens. Cliff, F. S. 1954. Snakes of the islands in the Finally, I wish to thank Patricia Blaney who Gulf of California, Mexico. Trans. San Diego typed large portions of the final dissertation Soc. Nat. Hist., 12:334-364. copy and Richard M. Blaney who is responsi- Cochran, Doris M. 1961. Type specimens of h\e for the reproduction of many of the photo- reptiles and amphibians in the United States graphs. National Museum. U.S. Natl. Mus. Bull., No. 220:1-291. Literature Cited Conant, Roger. 1965. Miscellaneous notes and comments on toads, lizards, and snakes from Allen, M. J. 1933. Report on a collection of Mexico. Amer. Mus. Novitates, No. 2205: amphibians and reptiles from Sonora, Mexico, 1-38. with the description of a new lizard. Occ. Conant, Roger, et al. 1956. Common names Pap. Mus. Zool. Univ. Michigan, No. 259: for North American amphibians and reptiles. 1-15. Copeia, 1956:1-26. Anderson, Paul. 1965. The reptiles of Missouri. Cook, Fannye A. 1954. Snakes of Mississippi. Univ. Missouri Press, Columbia. 330 pp. Mississippi Came and Fish Comm. Surv. AxTELL, Ralph W. 1959. Amphil)ians and Bull., 45 pp. reptiles of the Black Gap Wildlife Manage- Cope, Edward D. 1875. Check-list of North ment Area, Brewster County, Texas. South- American Batrachia and Reptilia. U. S. Natl. v/est. Nat., 4: 88-109. Mus. Bull, No. 1: 1-104. Bailey, Vernon. 1905. Report on the biologi- 1877. On some of the new cal survey collection of lizards and snakes and little known reptiles and fishes from the from Texas. N. Amer. Fauna 35:38-51. Austroriparian Region. Proc. Amer. Philos. 1913. Life zones and crop Soc, 17:63-68. zones of New Mexico. N. Amer. Fauna, 35: 1886. Thirteenth contribu- 1-100. tion to the herpetology of tropical America. Baird, Spencer F. 1859. Reptiles of the Proc. Amer. Philos. Soc, 23:271-287. boundary. United States and Mexican Bound- . 1890. Scientific results of ary Survey. 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Contributions to the herpetology of Sonora, Mexico: descriptions of new sul)- species of snakes (Micniroidcs ctiryxanthtts and Lanipropcltis ^ctulits) and miscellane- ous collecting notes. Amer. Midi. Nat., 54: 230-249. Volume 16, Number 3 '^'^3. coMp, ;o -^ May 6, 1970 I LIBRARY MAY 21 1970 Harvard UNiVERsjTY INTRODUCED GULF COAST MOLLUSCS DEE SAUNDERS DUNDEE Department of Biological Sciences, Louisiana State University in New Orleans, New Orleans, Louisiana 70122 p. 101 TYPE SPECIMENS OF FISHES IN THE TULANE UNIVERSITY COLLECTION WITH A BRIEF HISTORY OF THE COLLECTION ROYAL D. SUTTKUS Department of Biology, Tulane University, New Orleans, Louisiana 70118 p. ii6 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. 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Price per number (flat rate) : $1.50 Price per volume (flat rate) : $5.00 Meade Natural History Library Tulane University Riverside Research Laboratories Route 1, Box 46-B Belle Chasse, Louisiana 70037 Gerald E. Gunning, Editor Alfred E. Smalley, Associate Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Vickie L. Haddenhorst Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 16, Number 3 May 6, 1970 INTRODUCED GULF COAST MOLLUSCS DEE SAUNDERS DUNDEE Department of Biological Sciences, Louisiana State Univeisittj in New Orleans, New Orleans, Louisiana 70122 Abstract Of the ten species of molluscs which are known to have been introduced into the Gulf Coast area of the United States, four have been studied over a five year period; two of these, Brach/])aciui siiiiilaris ( Ferus- sac ) and Lamellaxis gracilis ( Hutton ) , were given concentrated attention; the other two, Riunina decollata (Linnaeus) and Gulella hicolor (Gould), while not studied as intensively, were used for com- parative puq^oses. Study of dispersal since introduction showed that these animals are still spreading. The means of dispersal ap- pears to be through man's activities re- lating to the transport of plants. Studies of biological requirements and tolerances by the snails indicate that they are well adapted to the types of habitats found in this area. Life history and ecological data reveal that further spread is likely. Introduction Some members of the gastropod fauna of the United States are not native. They have been introduced in various ways to our shores from other land masses. Such biological in- troductions often pose problems in the con- trol of the immigrant species, largely because we have no knowledge of rates, and in many cases the means, of the distribution or changes of ecological tolerances on the part of the introduced organisms. The goal of this study was to take a seg- ment of an introduced fauna and try to de- termine from whence it came, when and how it was introduced, and the rate and means of dispersal since being introduced. It was pro- posed also to study some of the ecological factors influencing the distribution of these introduced species, including a morphologi- cal study. Data gathered in this latter study will be published separately. The reported introduced fauna of the Gulf Coast study area (Houston, Texas to Mobile, Alabama: Figure 1 ), at the inception of this study numbered ten species. Frt)m these, four were selected for the purpose of study. They were chosen on the bases of availability and ease of raising in the laboratory. Two of them, Lamellaxis gracilis ( Hutton ) and Brady baena similaris ( Ferussac ) , were se- lected for intensive study while the others, Gulella bicolor (Gould) and Riimina de- collata ( Linnaeus ) , were examined more casually. The reason for limiting the study to these four was simply a matter of keeping the work involved within reasonable limits. Procedural Methods Effecting the proposed study resulted in dividing it into several phases: 1. Points of Origin; When and How In- troduced This phase of the project dealt largely with the correlation of past and present molluscan distributions with ship routes, cargoes, and currents. It involved literature searches and discus- sions with port authorities and ship- ping companies. It was necessary here to become acquainted and maintain contact with the members of the United States Department of Agricul- ture Quarantine Division. 2. Rate and Means of Dispersal This phase of the study consisted of making collections of these animals throughout the Gulf Coastal study area Editorial Committee for this Paper: Dr. Albert R. Mead, Department of Biological Sciences, University of Arizona, Tucson, Arizona 85721 Dr. Alfred E. Smalley, Department of Biology, Tulane University, New Orleans, Louisiana 70118 101 102 T/dane Studies in Zoology and Botany Vol. 16 \ • . . • • > • ) • • 1, •« n,. • / • J • • 1 • 0%: Figure 1. The study area. From Houston, Te.xas to Mobile, Alabama and northward to the level of the northern border of Louisiana. Dots indieate collection sites. to determine present distributions (Fig- ure 1) and the making of several fol- low-up collections to determine if fur- ther dispersal is occurring at present. Correlated with this phase of the study were visits to major United States col- lections to check further on distribu- tions. Determining means of dispersal proved to be a difficult task. Involved in this was the questioning of individ- uals such as nurserymen, gardeners, and so forth and it was not always easy to obtain accurate answers. Sometimes these individuals became suspicious that we might somehow be accusing them of wrong-doing in having these animals and they would then become defensive and not give complete an- swers. Most, however, were very help- ful. Careful surveys of all nurseries and greenhouses in New Orleans and Baton Rouge were made to determine what introduced species were there (Dundee and Watt, 1962; Hermann, Strickland and Dundee, 1965). Time limitations made it necessary to con- centrate detailed studies of this phase at the Port of New Orleans. 3. Habits of Introduced Molluscs It was also necessary to study in de- tail the habits of the molluscs both in field and laboratory. Laboratory colo- nies were begun in 1959 and from then through 1967, there were only brief periods when we were without colonies of these animals. Data on food habits, reproductive habits, lon- gevity, variability, ecological tolerances. and life histories, were gathered from these laboratory colonies. Colonies were easily maintained in the laboratory. Clear plastic containers of various sizes were used. Into each was put a layer of sand with a piece of paper toweling over part of it. The containers were kept damp with tap water in an air-conditioned rooni where the temperature ranged between 22- 25°C and the relative humidity near 40 percent. The significant humidities in the containers were much higher but no special check of them was made in routine daily care. One side of the room, which had an eastern exposure, contained large glass windows, hence the animals were regularly exposed to daylight. Food consisted of head lettuce leaves. The containers were cleaned and the toweling replaced at least every other day. Simultaneously, similar data were gathered from field stations for each of these species. In addition, marking of field individuals with nail polish gave some information on rates of movement. Acknowledgments This work was made possible through United States Public Health Service research grant GM-07194. I extend my sincere ap- preciation to that organization. I am also indebted to my various student assistants: Mrs. Patti Watt Hermann, Mr. Baldwin Shields Stutts, Mr. John Deris, and Mrs. Dian Hitt Sanders. All of them have been of much help. My husband. Dr. Harold A. Dundee, has accompanied me on many of the field trips and has helped make many of the collections. To him I owe sincere thanks. Findings 1. Points of Origin; When and How Intro- duced This discussion will be based entirely upon the Port of New Orleans. Of necessity, the discussion of introductions must be specula- tive since there is no record of the actual moment and method of introduction. The first recorded use of the port was in 1699 with the first settlements in the area of present-day New Orleans (Sinclair, 1942). No. 3 Introduced Gidj Coast Molluscs 103 Table 1. Point of origin of each species of mollusc and its distribution in the period 1699-1905. SPECIES ENDEMIC TO: DISTRIBUTION DURING PERIOD Bradijhaena .similciris (Ferrusac) Lumcllaxis gracilis (Hutton) (Uilclla J)icolor ( Could ) Ruiuina decoUata ( Linnaeus ) Southeast Asia Tropical Americas India, southeast Asia Mediterranean area: S. France, Italy, Sicily, Canary Islands Brazil, W. Indies, Hawaii, Mauritius Tropics of world; Mobile, 1885 ( Binney ) Tropics of world; W. Indies, Cuba, S. America In Bermuda and Havana ( Smith, 1912); in Philadelphia 1897 (Johnson, 1900) Thus we know that these species must have been introduced after that time. The first record of each of them in the New Orleans area is as follows: Brady baena similaris (Ferussac) 1939 (Harry, 1948) Lamelldxis gracilis (Hutton) 1905 (Pilsbry,'l946) Gulella bicolor (Gould) 1958 (Dundee, 1961) Rufz/iaa decollata (Linnaeus) 1906 (Piisbry, 1946) Gulella bicolor (Gould) was observed about 1940 by Dr. H. Harry (personal com- munication ) but was not reported. Know- ing when the port was first used and the first recorded date of the animals in the city sets the time of introduction of these species. If one now determines the point of origin of each of the species and the distribution at the turn of the 18th century as shown in Table 1, one then can compare those geo- graphical positions with points of origins of cargoes inbound to New Orleans ( Table 2 ) Table 2. Brief history of the Port of New Orleans. DATE IMPORTS ORIGIN OF IMPORTS 1699 1710 1710 1718 1720 1763-1776 1803 1806-1813 1812 1812-1862 1862 1862 1865 1871 1874 1899 Late 18()0's Provisions France Provisions, brandy, tobacco Cuba, Santa Domingo, Martinique, W. Florida Furs Ohio Valley FOUNDING OF NEW ORLEANS Slaves West Africa Powder, lead, clothing Great Britain Provisions, first coffee France, Spain, Brazil LOUISIANA PURCHASE— MANY PEOPLE MOVING IN Various items pirated from Spain Spanish ships and smuggled into the port FIRST REGULAR STEAMBOAT FROM NATCHEZ, MISSISSIPPI France Great Britain, CJuba and the islands Wines, brandies, cutlery, tools, China, cotton goods, sugar, rum, coffee, fruits, mahogany, rose- wood, silver, cochineal, pimento BLOCKADE OF THE PORT. No imports except cotton, sugar, and rice. QUARANTINE OF PORT AGAINST YELLOW FE\ER ALL CIVIL WAR RESTRICTIONS REMOXED . MANY NEW SHIPPING LINES ESTABLISHED FIRST RAILROAD Bananas Colon, Ecuador, Honduras, Guate- mala, Costa Pica Tropical fruits United Fruit Compan\- established Rubber, palm kernels, sisal, W. Coast Africa, Belgian Congo hardwoods 104 Tulane Studies in Zoology and Botany Vol. 16 Table 3. United States Depaitiiient of Agriculture interceptions of molluscs during 1959 and 1960 in New Orleans (Morris, 1959, 1960). GENUS CARGO POINT OF ORIGIN BulimuJtis corneus (Sowerby) Chiliim cf. fluminca (Maton) Cochlicclld hdibdia (Linnaeus) HcliccUa ( 4 spp. ) Limax nuirginatus Miiller Solaropsis- monile ( Rroderip ) Succinea sp. Tliehii piscina (Miiller) Zoiiitoidcs arhon'us (Say) Bananas Plant General cargo General cargo Cut flowers Bananas General cargo General cargo Orchid Ecuador Brazil France, N. Africa France, Greece, N. Africa Guatemala Ecuador N. Africa N. Africa, Italy, France Philippines and arrive at a logical conclusion as to ap- proximate time and means of introduction. These animals are assumed to have arrived by ship, since New Orleans was surrounded by a swampy wilderness divided by a network of waterways. Road and railroad travel were limited until much later (Civil War period ) . The first railroad travel was available in the 1870s. Even as late as the early 1800s over- land travel by means other than railroad was practically non-existent thus negating the likelihood that snails were brought in from Mexico and or Central America overland. Table 2 summarizes the history of the Port of New Orleans in relation to this discussion. Since 1865 many ships have included New Orleans on their regular itineraries. Some of those arriving from endemic areas for these snails carry cargoes which are likely ones for harboring such animals. Actual interceptions of molluscs by the Quarantine Division, United States Depart- ment of Agriculture in New Orleans in sev- eral recent years point out the potential for molluscan introductions. For example, in re- lation to this present problem, in I960 one of the interceptions at the Port of New Orleans was Lamellaxis ma//ritianus (Pfeiffer) (m an orchid from Venezuela (Morris, 1960). Table 3 summarizes several such intercep- tions during the years 1959, I960. Studies on other molluscs have also revealed that they too are arriving in this country on imported materials and it would seem that nursery stocks provide an ideal method of entry. In- spections will not reveal each single individ- ual which might be hidden on the plant parts or the ova which might be in the soil around the roots of the plants. It rapidly became obvious that these ani- mals are arriving through man's activities rather than being passively carried on float- ing vegetation or the like. Also, it is obvious that most of the introductions are accidental even though there have been cases where individuals have brought snails from another country to use as food items here. Many of these "foreign" snails can be bought alive in the markets of our country as food items (Figure 2). The question then becomes: what is to prevent them from crawling out of the container and establishing a colony? It is my belief that that is exactly how species such as Helix aspersa Miiller and others originally were established in the port area of New Orleans. 2. Rate of Dispersal All four of the migrants with which this paper is concerned are widely distributed in the warm areas of the world at the present time. Within the United States both Brady- haena similaris and Riimina decollata seem to be pushing northward as evidenced by our collections. Figure 3 shows the present known distribution of B. similaris while BM^jiyMpfY^ Ir^^^^H HBlinirTiiir* i *■' } ^m EmB^^^HH ^^Mk k 4 I'igure 2. Snails for sale. Iklix ci.spfrsa at 350, and Otala lactca at 750. No. 3 Intrnd/iccd G/dj Coast Molluscs 105 BRADYBAENA SIMILABIS Figure 3. Present distribution of Brachjbacna similaris in the United States. Figure 6 gives that of R. decollata. Gulella bicolor ( Figure 4 ) by contrast, still seems to be confined to the area where it was intro- duced— at least here in the Gulf Coast area. Lamellaxis ( Figure 5 ) , at first glance, would seem to be widely distributed in the eastern half of the United States; however, most of the northern records are greenhouse localities. Within the designated study area Brady- baena and Lamellaxis are both slowly widen- ing their distribution as shown by our re- peated collections of the area ( Figures 7 and 8). When this study began in 1959, B. similaris seemed to be limited to the New Orleans-to-Baton Rouge and Mobile areas. • GRACILIS ▲ CLAVULINUS ■ MICRA ^ MAURITANUS Figure 5. Present distribution of Lamellaxis in the United States. By 1961 we found it on the Mississippi Gulf Coast, spreading northeastward into south- western Mississippi and northward to Alex- andria, Louisiana. By 1962 it had been intro- duced to Hammond, Louisiana on the north and to Houma and Jeanerette on the south, and towards the mouth of the Mississippi River on the southeast. In 1964 it had moved into Thibodaux, Louisiana on the southeast. Complete further checks have not GULELLA BICOLOR Figure 4. Present distribution of Gtdella bi- color in the United States. Figure 6. Present distribution of Ihiniina (U colldta in the United States. 106 Tulane Studies in Zoology and Botany Vol. 16 0 I960 * 1961 % <962 ■ 1963 ^ 1964 Figure 7. Study area distribution of Brady- haena similaris showing dispersal over a 5-year period. Figure 8. Study area distribution of Lamel- laxis- gracilis showing dispersal over a 5-year period. Explanation as for Figure 7. been made since; however, plans are to sur- vey again soon. Lamellaxis gracilis distribution has also been carefully observed over a five year period (Figure 8). By 1959 and I960, L. gracilis seemed to be limited to New Orleans, Baton Rouge, Biloxi, Houston, and Beau- mont. In 1961 we found it spreading from the Biloxi locality into neighboring Gulfport, Moss Point, and Pass Christian in Mississippi. At the same time it was found in Picayune and Hattiesburg, Mississippi, probably origi- nating from New Orleans. The year 1962 found it established to the south of New Orleans in four localities and in one more to the north. By 1963 it had managed to ex- tend about 200 miles further north into Natchez, Mississippi and Natchitoches, Loui- siana. In 1964 it was still pushing westward from New Orleans along the lower half of the state. More recent findings show it in Pensacola, Florida on the east and Lake Charles, Louisiana and Galveston, Texas on the west. 3. Means of Dispersal There are two ways in which land molluscs are distributed: actively by their own move- ment, and passively by being transported by other organisms or the environment. Doubt- lessly most of the distribution of molluscs which we see today can be accounted for by the activity of the molluscs themselves over millions of years. However, with the numer- ous forms of transportation of modern man, many molluscs such as those involved in this work find themselves in areas far from their native habitats. Such introductions often pose severe problems in the control of intro- duced species. With the absence of natural population checks (predators, diseases, and the like), the immigrant is free to "run wild" biologically until a balance is again estab- lished in the ecosystem. The questions often asked are these. How are these organisms being transported? Why are we unable to stop introductions? An- swers, of course, vary with the organisms in question. It has become increasingly obvious during this study that the four mollusc spe- cies herein considered are being transported through the movements of plant materials. This is being further borne out by another study on introduced slugs now in progress. It was also found to be true in New York state ( Karlin and Naegele, I960). We first suspected this in noticing that these molluscs were always found in association with man's activities — in cultivated areas, near import centers, and so forth. Never have we found them away from man's activity areas even though habitats elsewhere appear to be satis- factory. If then we were correct that these snails were moving about as passengers on vegetation, where would be the most likely spots to find them? In gardens, public and private. In cemeteries. In nurseries and greenhouses. Private gardens were ruled out immediately as a study source because of the problems associated with entering such prop- erty. Checking public gardens and ceme- teries resulted in finding many new localities for the snails. If we could find the same snails in greenhouses, we could reasonably assiune that they got there with the plants ( as opposed to riding on bird feet, drifting on floating materials and so on). Such exam- inations did reveal the snails in many cases. No. 3 Introduced Ci/lj Coast Moll//scs 107 p? _x2z£izH_ lJL^ 12 10 s ..J«J1 =11 o HI 3 ■ lliff^ It also revealed other molluscs not previously reported from the area. Our next question was that of why it is that these snails are not discovered and killed. Many discussions with owners and operators of nurseries and greenhouses made us aware that there is a constant battle between man and snail. All owners and operators use some type of molluscacide and are constantly trying to prevent plant damage from these animals. It also became obvious that, despite the fact that they wish to rid themselves of such pests, they are not thorough enough. We were able to find many molluscs under the tables where nothing economically im- portant was being grown and where they are not using the chemicals. It was difficult to find snails on the about-to-be-sold plants, however. How then, were these snails being distributed? Our studies have revealed that they spread largely through transport of snail ova which have been deposited in the soil around the plants. >- z o _l o o , ■ J n F ^1 < a: < z < -I ,allL 12- .-_!. 18 16' _-_t jllL ooooooooooo HEIGHT IN MV 1= W/0 OVA ^ = WITH OVA 4. Future Dispersal If, in the five years of studies, these snails have spread considerably through much of the study area, what then of the future? How far can they go? Baker (1958: 143) sum- marizes it well: "In time, dispersal means that each species would occur everywhere unless it be stopped by barriers . . . " Basically there are two types of barriers which prevent an organism from spreading everywhere: physical (geographical) and ecological. Since we have already established in this particular case that physical barriers are easily spanned with the aid of man, the immediate question becomes: Are there any ecological barriers which will stop the spread of these animals? Can they exist wherever man takes them? To answer these questions it was first necessary to determine something of the biology of the animals. A. Life History: Lamellaxis gracilis. In Figure 9 one can see that, in the New Orleans area, the reproductive season is initi- ated in mid-March at which time ova may be seen through the shell. This situation pre- vails to late May at which time the ova are deposited. Young of various ages are present Figure 9. VkU colon) , Lamellaxis gracilis. 108 Tidane Studies in Zoology and Botany Vol. 16 Figure 10. Ova in soil, Larnellaxis gracilis. from early June through the summer ( Figure 9). By about October they have grown to approximately 4-5 mm in height. Early October is another period of ova deposition followed by some young overwintering in the 3 mm stage. Laboratory colonies maintained for over a year tend to behave in a somewhat different manner in that March to June is a period of ova production and is followed by an interval of no ova nor young and then by another peak of production of ova in late summer followed by constant low ova production for the rest of the year. This is most likely the result of the colony being in a more or less constant temperature-humidity atmosphere (temperatures from 22-25 ""C and relative humidities constantly in the 40 percent range ) . The maximum number of ova ever seen in one snail was 10; the average number per gravid snail was 5 seen at one time. How- ever, based on laboratory experiments, the average number of ova deposited per snail per breeding season is 79. In lab colonies 55 percent of the ova hatched. These ova ( Figure 10) of Larnellaxis gracilis are almost spherical, white, have "hard shells," and are deposited singly in the surface of the soil, but not covered by soil. There appears to be no effort at "nest building"; the ova are simply placed individually in the soil; usually several ova will be found together as a clutch. Figure 11. Laboratory, Larnellaxis gracilis. Colony liegun witli 21 indiN'iduals and all ova and/or young removed from it regularly. _l < O > a: m ■s. 13 >- Z O o o m < I en < a: CD X < UJ < No. 3 Introduced Gulf Coast Molluscs 109 MOVEMENTS OF LAMELLAXIS Figure 12. Movements of marked individuals, Latucllaxis gracilis. These ova are small enough ( 0.5-0.9 mm in diameter ) to be easily overlooked by indi- viduals transporting plant materials. Incubation time in nature is approximately 10 days. However, on 2 occasions in the lab- oratory incubation was completed in 7 days. Larnellaxis micra, by contrast, had an incuba- tion time of 15 days in the laboratory. Its ova are similar but smaller. Larnellaxis gracilis growth rate is approxi- mately 1 mm month. Those young which hatch in mid-June are in the 5 mm class by late October to early November. This rapid growth rate then seems to slow with a con- tinued constant, gradual growth until death. Figure 1 1 shows a laboratory colony which was begun with 21 adults of various sizes. Constant removal of all ova and 'or young immediately upon discovery kept this colony intact for over a year. During that time 10 individuals died and the remaining 11 con- tinued growth as shown on the histogram. Adult Larnellaxis gracilis ( as evidenced by the initiation of reproductive cycles) range in size from 4—14 mm in height and have whorl numbers between 5.5-10. Individuals exceeding 12 mm in height are rare. Larnel- laxis micra adults range in size between 4.5 and 7.2 mm in height. The life span of an individual L. gracilis appears to be approximately one year with some apparently surviving through most of the second year. These observations need substantiation since it was impossible to ob- tain recapture of marked individuals after a lengthy period. B. Habitat and Habits: Larnellaxis gracilis. These snails are found in areas of man's activities — in particular, in areas where man has been active and then has allowed the area to stand idle for a period. A prerequisite is cover of some sort — boards, bricks, leaves, or other objects which can provide shelter. The snails are always under these objects dur- ing periods of dryness. After rains they are easily available crawling about on the sur- face. At no time are they more than 1.3 cm or so above ground. As cooler weather ar- rives, Larnellaxis gracilis descends into the soil. During one period of -5 C, the animals were found 7.6 cm down in the soil. During a January period when the soil temperature stayed close to 8°C, the snails, despite heavy no Tidiwe Studies in Zoology and Botany Vol. 16 OCTOBER T Q UJ h- U UJ o u <0 < o > o u. o q: UJ CD AUGUST 4 SEPTEMBER 1 ■ ■ ■ I I 1 I I I I 1 I 1 I T I T I I r r T I I T I I I I I -r 1 OO'O^OtnO'J'OtJiOUiOtn OcjC'iJ»0 "-"omOtnOui o"*© FIELD BRADYBAENA Figure 13. Field colony, Bradybaciia simi- laris. rains, remained from 2.5-7.6 cm below the surface. Even at these depths individuals are grouped together; for example, in a 4.5 X 10 cm area 27 individuals were found 1.3 cm down in the soil. Under the circumstances described in this paragraph the individuals are lying dormant (hibernation?). Feeding experiments were run in the labo- ratory. Individuals were offered plant parts of each of the species of plants found in the habitat. Results indicated that they will feed readily on leaves of Laportea canadense, Pani- cum sp., and Crepis sp. Other plants from the habitat were not acceptable. Karlin's (1956) assumption was that since Lamel- laxis and two other species were never found more than an inch above soil surface, they were feeding "only on plant matter in the mulch. . ." The observation of the closeness to the soil is in accordance with our findings; however, it seems likely that these snails do use the leaf matter described above in their normal living but probably rely on those leaves which drop off or are low down. In the laboratory any of the species of Lamel- laxis seem to survive for long periods on nothing more than lettuce leaves. Repeated marking experiments all met with failure. One is shown in Figure 12. Here 6 snails were marked differentially with nail polish and released as indicated on Octo- ber 1. One week later we were unable to find 2 of them ( B, D ) . By a month and a half later (November 18) the other 4 were in the positions shown. With the advent of colder weather shortly thereafter, we were unable to get further recaptures of these. It is believed, however, that enough other re- captures were made — albeit not after lengthy periods — to state tentatively that L. gracilis seems to remain within 10 ft or less of its point of release; whether this is true during the entire life is unknown. C. Life History: Brady baena similaris. In the New Orleans area B. similaris de- posits its ova in January and by early Febru- ary many young snails are present ( Figure 13). The older members of the colony ap- pear to die off soon after the winter season as shown in Figure 13. Through spring and summer the young are growing and a middle- aged group is present. Again in late July there is a brief period of egg deposition so that by mid-August to early September there are again a few young present in the popula- tion. In this area these young reach a subaduk stage before cooler weather occurs. Just as with Lamellaxis the laboratory colonies con- tinue producing ova and young throughout the year in small numbers. Bradybaena ova (Figure 14) by contrast with those of Lamellaxis, are clear and gelati- nous when first deposited. They are fre- quently stuck together ( probably acciden- tally ) and always laid as a cluster. The adult usually makes a concavity in which the ova are placed; they are not covered with soil. Each ovum is spherical and measures ap- proximately 2 mm in diameter. In laboratory colonies the average number of ova per snail per breeding season was 38. Incubation time in the lab was 1 1 days and in the field it ranged from 10-15 days. The smallest Bradybaena similaris at hatch- ing measure 1 mm in height, 1.75 mm in No. 3 Introduced Gulf Coast Molluscs Figuii.' 14. C)\a in soil, Hnidijlxnitd .siinildiis. diameter, and have 1.75 whorls. Six monchs later most of these had reached 3.25 mm in height, 5.25 mm in diameter, had 3.5 whorls, were sexually mature and reproducing. All newly hatched young were without color bands which are characteristic of 91.1% of the population here. The bands began to show by the time the snail had reached 2 mm diameter and 1.5 mm in height. Of the 1566 Bradybaena similaris checked, 3.9% had no bands at all in the adult and 6% showed only very faint banding. The remainder have a characteristic light yellowish shell with a brown band. This stands in contrast to the same species in Japan ( Komai and Emura, 1955 ) where there are 4 distinct genetic types: the 2 mentioned above plus brown forms with and without bands. Why these introduced forms lack the brown phases is not known at this time. Adidt Bradybaena similaris in this area also differ much in size from those reported by Komai and Emura ( 1955 ). They report them to be "18 mm in diameter and 12 mm in height at most of its size." The tallest Bradybaena we have recorded in many mea- surements is 1 1 mm and the one of greatest diameter was 15.5 mm. These are rare. This smaller size, plus the absence of the two color phases, suggests that some factor in the habitat, present in Japan but missing here, is exerting a considerable influence on these snails. We plan to do some follow-up work on this phase. D. Habitat and Habits: Bradybaena simi- laris. It has been suggested that this species is closely associated with sugar cane ( Pilsbry, Hyatt and Cook, 1912). While we have found it in close proximity to sugar cane fields here, it is not necessarily associated with them. Many of the colonies are found living in various types of tall grasses in \acaiit lots, along railroad tracks, and so forth. Others, however, have become estab- lished in gardens. Sugar cane was introduced from Santo Domingo to this area between 1771 and 1775 in Jesuit Bend, a community 15 miles south of New Orleans. Since B. siDuLiris was known from the West Indies at that time (Table 1 ), it is quite probable that it was introduced here with the sugar cane. In feeding experiments we were unable to get B. sif?iilaris to eat anything but mono- cots and, in particular, the grass, Panicum brachyanthus and a lily, Liliuni canadense. All of the plants from their normal habitats were offered. These snails are much less secretive than Lamellaxis. They are just as likely to be found out in the open ( either active or dor- mant ) as under objects. Occasionally they are seen a few feet high on vegetation or on the sides of vertical objects. Usually they are not found over 15 cm above the ground on the plants. Most often they are seen at night crawling about on grass which previ- ously has fallen down and dried. Often hun- dreds of them can be seen in this type of situation. Marking experiments in 3 different locali- ties revealed that individuals move little at any time. The maximum distance traveled away from one point of release by any one individual was just under 3 m in a 2 month period. This, of course, is not taking into consideration how much movement occurred within the 3 m radius during that j>eriod by this individual. Marking was done with nail polish; this seems to lead to an early death in the snails. E. Ecological Tolerances. ( 1 ) Temperature: An experimental de- vice similar to, and patterned after, that used by Dainton ( 1954) was used to determine temperature tolerances of these snails. This device consists of a trough in which a tem- perature gradient can be maintained. Dif- ferent ranges can be had by using different water temperatures at the inputs. Snails are introduced throughout the length of the 112 Tulane Studies in Zoology and Botany Vol. 16 Q. X O m 20 I 9H le I 74 I 6 I 5- 14- 13 12 I I 10 9 8- 7- 6 5 4 31 2 I ■ NUMBERS INDICATE INDIVIDUALS SELECTING THAT TEMPERATUREI 2 4 2 21 6 3 I 2 2| 4 4 X = NUMBER KILLED JE I 4 2 3 2' :i] tiz 3 3 5 I fc 4 3 4 2 I I ,1 2 3 n — I — ] — I — I — I — ' — I — ^-—r — r — I — I : 1 — ' 1 1 — 1 ' — I — I — 1 [— rv — ojro— ou3(D->j(jim.t»OJro— OtDOD-sicnai^wrxj ,2,2: 3] -| — I 1 — r- Qt£)CD-Nlai ^UJfNJ uj 03 -J a> ui -t DEGREES CENTIGRADE 36 - 15 C TOLERANCE LAMELLAXIS GRACILIS TEMPERATURE SELECTION Figure 15. Temperature selections, Lamellaxis gracilis. trough and allowed a period in which to select the temperature most suitable for them. The results are recorded after 30 minutes. After repeated trials for each species, during which the temperature ranges were varied (Figure 15) so as to include all reasonably possible temperatures, it was found that the range of tolerance for La/nellaxis gracilis is 15-37°C while that for Bradybaena similaris is slightly greater: 12-38°C. Interpretations of these data must be made in conjunction with other factors. If one were to use only these tem- perature data, one would be forced to con- clude that these snails can live nowhere in the United States except Florida; as has al- ready been shown ( Figures 3 and 5 ) the snails do live elsewhere. If one plots upon the present distribution picture the average minimum temperature for January, one finds that these snails, which, according to our ex- perimental data can tolerate no less than 12°C, are living in areas where the average January minimum is 6°C. Here again, it is of interest to note that Komai and Emura (1955) found that -7°C for an hour or 2 killed nearly all adults; perhaps these snails are able to tolerate much lower temperatures in their native homeland. When one con- siders that these snails protect themselves from the cold as described above, then it is not t(X) surprising to find them living in colder areas than the laboratory experiments indicate are possible. During the period in which we were recording soil temperatures, the lowest reading was a December one of -2°C at a 3 inch soil depth. The snails at this level were aggregated and they had their epiphragms in position, but many of them were dead. This was true for both Lamellaxis and Bradybaena. The upper limits of tolerance of these snails are such that they should be able to tolerate temperatures anywhere in the United States since the highest average July tempera- ture falls easily within their range of toler- ance. The maximum could be withstood through the use of shelter. (2) Desiccation: Experiments consisted of merely placing the snails in dry bowls (one per bowl) at room temperature (23- 29°C) and humidity (407r) and examining them at intervals to see if they were still alive. It was found that dead ones could be easily identified by shining a narrow beam from a spot lamp through the shell and look- ing to see if the animal was intact. Surprising differences in tolerances to desiccation were revealed in our experiments. In several hun- dred trials, the time which Bradybaena was able to remain alive ranged from 30—217 days with an average time of death being 122 days, or approximately 4 months. In contrast, the maximum time which Lamellaxis was able to remain alive was 121 hours, or ap- proximately 5 days. To evaluate these data one must keep in mind that the 40% relative No. 3 Introduced G/zlf Coast Molluscs 113 Table 4. Molluscan associates of the intioducfcl species. SPECIES STATION STATION STATION STATION 12 3 4 Bradt/haena sim ilaris CucUii l)i<-()J()r Rum ilia dccollata LamcUaxis gracilis Hcliciua orlnculata Poli/fli/ra fcxa.siana Rcthulla iiidciitata Haicaiia iniiii.scula Ptipoidcs iiKir^inatiis Poliifiyrd sept em volva Pratieolella ^riseola Me.sodon pcri^rapfri.s Stroljilops texa.siana Dewceras laeve X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X humidity found in the lab during these ex- periments is low in contrast to that in the environment in this Gulf Coast area and the microenvironment of these snails. Thus, the animals should be able to withstand periods of drought longer in nature than they can in the laboratory. The fact that 3 of the 4 spe- cies treated here can survive adverse con- ditions for long periods of time through estivation is doubtlessly of high survival value. It is of interest that Lamelh/xis gracilis. despite the fact that it is widely distributed over the world now, does not have this ability. Obviously, a combination of other factors has overcome this one factor which would appear to be a distinct handicap to an introduced species. Similar experiments with Ri/mina decol- lata and G/della hicolor revealed that under the circumstances described some individuals of both species are able to survive at least 7 months. One G/della was still alive at the end of 214 days. Rascop (I960) found that relative humidity below 159^ and a tempera- ture above 85°F (29°C) caused Rumina to estivate. Further, Rascop found that snails estivating at room temperature for 8 months lost an average of 55.56% of their body weight. (3) Soil pH: Throughout our studies we found pH ranges at the various study stations of 6.4-7.1 for both Lafuellaxis gracilis and Brady baena similaris. Karlin (1956) con- cluded that pH could not be considered a critical factor in greenhouses since the mean pH where L. gracilis and others lived was 5.9. He pointed out that while it is well- known that snails are scarce on acid soils, other factors such as the addition of gypsum and the presence of other minerals compen- sated for the low pH. (4) Moisture: None of these 4 snails is associated with visible moisture in the form of swampy or marshy conditions. All live in well-drained areas. Among the 4, they could be ranked on the basis of 5 years of obser- vation both in the field and in the lab, ac- cording to their moisture tolerances (from wettest to driest ) : La???ellaxis, Bradyhaena, G/della. R/iviina. Observations lead us to believe that moisture, in the form of relative humidity, is of far more direct importance to the land mollusc than standing or falling water. Relative humidities of 857^^ and up are not uncommon in this area even in the day. A dry day would have a humidity in the 40% range. From this standpoint, then, conditions in this Gulf Coast area seem to be ideal for these snails. (5) Relative Population Densities: L. gracilis and B. similaris are, in the habi- tats described, extremely abundant. One can easily collect hundreds of specimens within half an hour. The habitats, however, are fre- quently widely separated so that this heavy abundance actually forms a spotty type of distribution. Some populations were large enough that we were able to remove 50 or more individuals every two weeks without any noticeable change in the numbers pres- ent. In contrast, G/della bicolor and R/imina decollata are abundant nowhere in this area. The distribution is not only spotty but within a given habitat one would be likely to find no more than 10 individuals in the half-hour period. 114 Tidane Studies in Zoology and Botany Vol. 16 (6 ) Associates: Table 4 reveals that there are no associates which are always found with a given introduced species. The most common joint occurrence is that of the two introduced species, B. siniilaris and L. gracilis. Apparently no significance may be attached to the presence of any of the molluscan asso- ciates of these introduced snails. Other associates such as ants (Formicidae) do seem to be extremely significant to the well-being of these snails, however. Ants are very common in this entire study area. We observed repeatedly that whenever ants in- vaded a habitat such as the area beneath a large boulder it would not be long before the snails would all be dead. We do not know the reason. We did run several simple ex- periments of enclosing the snails and the ants in gallon jars together and again the results were the same — the snails soon died ( controls did not ) . This, perhaps, might be investigated further with a view towards bio- logical control if such is ever needed. 5. Economic Considerations Fortunately, at the present time, none of the introduced molluscs in the Gulf Coast area have become severe pests under natural conditions. Even though they are a problem in greenhouses and nurseries, this is a differ- ent matter. B. similaris is an annoyance in a few flower gardens, but no more so than endemic species such as Mesodon thyroidiis. In any event, no serious damages to crops nor any nuisance from numbers of snails such as was the case with Acbatina jidica (Mead, 1961 ) has occurred in this area. Discussion We can, after examining the foregoing data, return to the question posed in the be- ginning of section 4 of this paper: Can these snails (in particular, B. siinilaris and L. gracilis) exist wherever man takes them? The general answer, of course, is no. There are many areas, however, where they do not now exist but where they could easily sur- vive. There is no cause to think that they will not spread further northward, eastward, and westward. They should be able to live throughout the lower half of the United States from southern Pennsylvania, the lower half of Ohio, Indiana, and Illinois, southern Missouri and Kansas, southeastern Colorado and along the entire west coast. The average winter minimum temperatures of this area are no less than -2°C (28.4°F). Since we are considering only average temperatures the snails in the northernmost part of this area would have to remain confined to shel- tered type habitats and, therefore, should be much more spotty in distribution than else- where. Drier parts of this range, namely in west Texas, New Mexico, Arizona, and southern California should act as barriers to the gen- eral spread in that direction. However, again, cultivated areas should provide habitats which have adequate moisture to maintain high enough relative humidites in the micro- habitats, especially for small snails such as Lamellaxis. Through follow-ups on this study we will know definitely whether, by a brief survey of such factors as have been dealt with in this paper, one would be able to predict the future dispersal of an introduced land mol- lusc. It seems now that such will be the case. It appears that most of the introduced snails which do become established do not become agricultural pests. Of the 10 species which are known to have been introduced in the New Orleans area at the inception of this study, 4 have failed to become established, 4, as shown in this paper, are still spreading, and 2 other species are also established and spreading. None of these has yet become a real pest to agriculture or horticulture. It appears also that preventing the en- trance of these foreign invaders into this country is impossible. The United States Department of Agriculture is doing an ad- mirable job of keeping down the numbers entering but even they are not preventing the entrance of many. As long as we import materials we will accidentally import snails. One of the best sources of foreign land mol- luscs in this country is the nursery. It is ideal for their reception, maintenance, and distribu- tion. The key to whether or not one of these immigrants will become a severe pest seems to lie in its food habits and its reproductive potential, assuming of course that it is able to find a suitable habitat. It appears that if an investigation of a species' reproductive and food habits were undertaken as soon as an introduced land mollusc is discovered, it should be easily possible to predict whether the immigrant would become a severe agri- No. 3 hitrodi/ced Gidj Coast Molluscs 115 cultural pest; if so, immediate control mea- sures could be instituted. Perhaps such a program could be under- taken by the United States Department of Agriculture through a program which would encourage malacologists and other interested biologists to be on the alert for new intro- ductions in their areas. If and when such is found the ideal situation would be for the United States Department of Agriculture to request that a study be made by a malacolo- gist, supported by U.S.D.A., to determine if the immigrant is likely to become a pest. If chances are good, then control action could be initiated promptly; if not, there would no longer be a need for constant vigil at our ports for that species. Literature Cited dispt Baker, H. B. 1958. Land snail dispersal Nautilus 71(4): 141-148. BiNNEY, A. 1885. Nhmual of Amt'rican Land Shells. U. S. Nat. Mus. Bull. 28:1-426. BuRCH, J. B. 1960. Some snails and slugs of (juarantine significance to the LInited States. Sterkiana N.(2): 13-56. Dainton, B. H. 1954. The activity of slugs. J. E.xp. Biol. 31:165-197. Dundee, D. S., and P. Watt. 1961. Louisiana land snails, witli new records. Nautilus 75(2): 79-82. . 1962. New Orleans greenhouse gas- tropods with a list of distributions of some other southern snails. Proc. Louisiana Acad. Sci. 25:47-49. Harry, H. W. 1948. Notes on foreign snails of Louisiana. Nautilus 62(l):20-24. Hermann, P. W., B. C. Stricki^no, and D. S. Dundee. 1965. Baton Rouge greenhouse gastropods. Nautilus 78( 3) : 131-133. lon.NsoN, C. W. 1900. Some notes on Huiniin (IccoIIafa. Nautilus 13(10):117. Kahlin, E. J. 1956. Notes on the ec()log>' of Zoiiifoides arhorcus (Say), Opt'as- fiuiuihim (Pfeiffer) and Lantcllaxis firacilis (Hutton) in greenhouses. Amer. Midi. Nat. 55(1):121- 125. Karlin, E. J., and J. A. Naecele. 1960. Biol- ogy of the Mollusca of greenhouses in New York State. Cornell Univ. Agr. Exp. Sta., Memoir 372:1-35. KoMAi, K., and S. Emura. 1955. A study of population genetics on the polymorpliic land snail Bradt/haena sinuhiris. Evolution 9:400- 418. Mead, A. R. 1961. Tlie giant African snail: a problem in economic malacology. Univ. Chicago Press, 257 pp. Morris, S. R. 1959. Plant pe.st interceptions at Louisiana ports. In: Insect Conditions in Louisiana. Entom. Res. Dcpt., Louisiana State Univ. Agr. E.xp. Sta., pp. 40-46. . 1960. Interceptions at Louisiana ports in 1960. In: Insect Conditions in Louisiana. Entom. Res. Dept., Louisiana State LJniv. Agr. Exp. Sta., pp. 41-46. PiLSBRY, H. A. 1946. Land mollusca of North America (north of Mexico). Volume 2, 520 pp. PiLSBRY, H. A., H. Hyatt, and \L Cook. 1912. Manual of Conchology, 2:22, 25. Rasc:op, A. M. 1960. The biology of Riiniina decoUaiu (Linnaeus), Pulmonata: Achatinidac. Unpubl. Master's thesis, L^niv. Ariz., pp. 1-65. Sinclair, H. 1942. The port of New Orleans. Doubleday, Doran (!sc Co., Cardcn City, N. Y., 335 pp. Smith, H. H. 1912. Ruiiiina dccolUita in Mo- bile and New Orleans. Nautilus 26(1 ):4. TYPE SPECIMENS OF FISHES IN THE TULANE UNIVERSITY COLLECTION WITH A BRIEF HISTORY OF THE COLLECTION^' - ROYAL D. SUTTKUS Department of Biology, Tulane University, New Orleans, Louisiana 70118 Introduction A brief history of the Tulane University fish collection and a list of type specimens are given in this paper. The publication of the latter is recognized as part of the Uni- versity's responsilibility following the recom- mendations of the International Code of Zoo- logical Nomenclature (Recommendation 72 D. Institutional responsibility) 1961. For- merly, some paratypes were housed elsewhere (as noted) but were acquired by Tulane on a gift basis. All types are grouped in a sepa- rate room apart from the rest of the general collection. Jars containing holotypes are en- circled with red paint and those containing paratypes are similarly marked with blue paint. The types as well as the general col- lection specimens are arranged by families in a phylogenetic sequence following Berg's (1947) classification system. Within the family the genera and species are arranged alphabetically. Each label bears a family number which has been derived by a renum- bering of Berg's families of fishes which con- tain living representatives, that is, families with only fossil forms were excluded because fossil fish material is kept separate from extant material. History of the Tulane University Fish Collection During the latter half of the last century Tulane University had an active exhibit museum which included an array of natural history items as well as art items, minerals, and so forth. In the early 1950's when the old museum was disbanded there were only a few fish specimens remaining and these were suspended by thread in formalin. None had labels and so the several dozen specimens were discarded. Two dry-mounted fish speci- ^ Contribution Number 1 from the Laboratory of Systematic and Environmental Biology, En- vironmental Science Center, Tulane University. - Published by Tulane University in fulfill- ment of an institutional responsibility recom- mended by the International Commission on Zoological Nomenclature, with partial financial support from the Louisiana Research Founda- tion. mens, a cotton stuffed Acipenser fidvescens which has "Niagara Falls, July 1881" printed on its ventral side a partial skin of a Rachy- centron canadus which has "New Orleans, June 3, 1894" written on the exposed part of the artificial wooden body are the only specimens retained from the former Tulane Museum collection of fishes. The author started the modern Tulane University fish collection in the fall of 1950. However, it was not until July 11, 1951, that the first entry was made in a permanent cata- log. The collection has grown steadily and at present, December 31, 1969, there are 60,030 catalog entries which total 1,849,315 specimens. Some of the earlier series, 1951 to about 1955, were stored in 707^ ethanol, but the bulk of the specimens have been stored in 40% isopropyl alcohol, which is the currently used storage fluid. Of the two percentages of isopropyl alcohol available commercially (99% and 91%) 99% is used for dilution to 40%^^ with ordinary tap water. The fish skeletal collection totals 549 specimens, most of which are complete skele- tons. The dry fish skeletal material and the cleared and alizarin-stained, glycerin stored specimens are cataloged separately from the above mentioned alcohol stored material. Moreover, the 953 x-ray negatives are cata- loged separately on 4 X 6 file cards. The 953 x-rays represent a total of 12,042 speci- mens. The Tulane University fish collection is not a typological or local reference collection. Geographically the Tulane collection has ex- cellent representation of southeastern, south- ern and southwestern United States fresh water fishes and good representation of cen- tral United States, Mexican, Central Ameri- can (particularly Costa Rica) fresh water fishes and marine fishes from the Gulf of Mexico and Caribbean Sea. During the entire two decade period the author has been the curator of the fish col- lection. During the first decade of history a number of students helped in curating the collection, in particular. Sue Whitsell (Fin- german), Myrna Andersson (Wilkens) and 116 No. 3 Type Specimens of Fishes at Tiilane 117 Bangalore I. Sundararaj. In 1963 John S. Ramsey became the first volunteer student curator, followed by Clyde D. Barbour and Glenn H. Clemmer. Ramsey organized the type material; sorted, identified and cata- loged a vast amount of material; studied and identified most of the unidentified cataloged material that had accumulated up to that time and took care of most of the loan re- quests during the 1963 to 1965 period. Dr. D. E. Copeland (then Chairman of the Zoology Department) was instrumental in obtaining the first curatorial staff appoint- ment for the fish collection. Mr. Albert Harris was appointed as assistant to the curator of fishes in 1965. Upon his resigna- tion in 1966, Mr. Frank Thomas became the assistant to the curator and has continued in this position to the present time. Without the staff appointment the ever increasing requests for loans would have been an overburdening task for the curator. Needless to say the growth of the collection and the production of publications would have been greatly curtailed without the assist- ant. Too, during the two decade period parts of the collection have been moved several times from one building to another on dif- ferent sides of the campus and at the present time we are in the process of moving the en- tire collection to the Environmental Science Center which is 15 miles away on the west bank of the Mississippi River. List of Type Specimens of Fishes in the TuLANE University Laboratory of Systematic and Environmental Biology* Clupeidae Brevoortia gunteri Hildebrand, 1948, Smith- sonian Misc. Coll. 107(18) :l-39, 9 figs. Paratype: TU10792 (formerly United States National Museum No. 129802, sent to Tulane University as gift exchange on Janu- ary 5, 1954), Texas, Aransas County, Aransas Bay, collected by Gordon Gunter. Alepisauridae Alepisaiirns brevirostris Gibbs I960, Brevi- ora, Mus. Comp. ZooL, ( 123 ): 1-14. * The formal opening of the Laboratory of Systematic and Environmental Biology will hv announced during the latter part of 1970 or early 1971. Paratype: TU 22990, Atlantic Ocean, lati- tude 37^ 44' north, longitude 65 '^ 42' west, 1957, collected by M V Delaware. Cyprinidae Algansea monticola Barbour and Contreras, 1968, Proc. Biol. Soc. Wash., 81 : 101-108. Holotype: TU 40869, from Mexico, Za- catecas, Rio Juchipila near town of Jalpa, at Highway 70 bridge, one mile east of the intersection with Highway 41, 30 June 1963, CDB 63-27, collected by Clyde B. Barbour and Salvador Contreras. Paratypes: TU3061I, 49 specimens from same collection as holotype; TU30608, 76 specimens, from Mexico, Zacatecas, Rio Juchipila at Mai Paso (a little walled town) about 16 miles east of Ciudad Garcia Salinas, 28 June 1963, CDB 63-23, collected by Bar- bour and Contreras; TU 30609, 131 speci- mens, from Mexico, Zacatecas, tributary to Rio Juchipila, about 5 miles south of Vil- lanueva on Highway 41, 28 June 1963, CDB 63-24, collected by Barbour and Contreras; TU 30610, 18 specimens from Mexico, Za- catecas, Rio Juchipila at Tabasco, upstream of bridge at south end of town, 29 June 1963, CDB 63-25, collected by Barbour and Contreras. Nocomis effiisiis Lachner and Jenkins 1967, Copeia, 1967(3):557-580. Paratypes: TU 19488, 2 specimens from Tennessee, Rutherford County, Stones River, 6.5 miles north of Murfreesboro, Highway 231, 7 September 1958, RDS 2758, collected by Suttkus and Myrna Andersson; TU 32838, 5 specimens from Tennessee, Fentress County, tributary to Wolf River, 9.8 miles north of Jamestown, Highway 127, 3 June 1964, RDS 3474, collected by Suttkus and John S. Ram- sey; TU 33115, 2 specimens from Tennessee, Cheatham County, Harpeth River, 8 miles east of White Bluff, Highway 70, 15 July 1964, JSR 64-60, collected by Ramsey and Dan W. Walton; TU 33125, 11 specimens from Tennessee, Williamson County, South Harpeth River, 4.2 miles east of Fairview, Highway 96, 15 July 1964, JSR 64-61, coL lected by Ramsey and Walton; TU 33171, 21 specimens from Tennessee, Rutherford County, West Fork Stones River at west edge of Murfreesboro, Highway 96, 15 July 118 Tulane Studies in Zoology and Botany Vol. 16 1964, JSR 64-63, collected by Ramsey and Walton; TU 33194, 1 specimen from Ten- nessee, Rutherford County, Stones River at south limit of Walterhill, Highway 231, 16 July 1964, JSR 64-64, collected by Ramsey and Walton. Notropis asperijrons Suttkus and Raney, 1955, Tulane Stud. Zool., 3(1): 1-33. Paratypes; TU 2974, 22 specimens from Alabama, Wilcox County, Gravel Creek trib- utary to Pursley Creek 6.3 miles south of Camden, Highway 11, 3 June 1951, RDS 2140, collected by Royal D. Suttkus and Charles D. Hancock; TU 3063, 5 specimens from Alabama, Wilcox County, tributary to Pursley Creek, 1.8 miles east of Camden, Highway 10, 3 June 1951, RDS 2138, col- lected by Suttkus and Hancock; TU 3426, 8 specimens from Alabama, Wilcox County, Pursley Creek, tributary to Alabama River, 3.4 miles southwest of Camden, Highway 11, 3 June 1951, RDS 2139, collected by Suttkus and Hancock; TU 4251, 28 specimens from Alabama, Shelby County, tributary to Waxa- hatchee Creek, 4.7 miles southwest of Colum- biana on Highway 25, 15 June 1952, RHG 356, collected by Robert H. Gibbs, Jr. and Philip P. Caswell. Notropis bailey i Suttkus and Raney, 1955, Tulane Stud. Zool., 2(5) : 69-86. Paratypes: TU 2637, 29 specimens from Alabama, Monroe County, tributary to Ala- bama River, 17.3 miles south of Camden, Highway 11, 3 June 1951, RDS 2141, col- lected by Suttkus and Hancock; TU 2965, 4 specimens from Alabama, Wilcox County, Gravel Creek, tributary to Pursley Creek, 6.3 miles south of Camden, Highway 11, 3 June 1951, RDS 2140, collected by Suttkus and Hancock; TU 3066, 41 specimens from Ala- bama, Wilcox County, tributary to Pursley Creek, 1.8 miles east of Camden, Highway 10, 3 June 1951, RDS 2138, collected by Suttkus and Hancock; TU 3195, 4 specimens from Alabama, Butler County, Pine Barren Creek, 2.5 miles south of Forest Home, 2 June 1951, RDS 2136, collected by Suttkus and Hancock; TU 3430, 75 specimens from Alabama, Wilcox County, Pursley Creek, tributary to Alabama River, 3.4 miles south- west of Camden, Highway 11, 3 June 1951, RDS 2139, collected by Suttkus and Hancock. Notropis edivardraneyi Suttkus and Clem- mer, 1968, Tulane Stud. Zool. and Bot., 15(l):18-39. Holotype: TU 49485, from Alabama, Wil- cox County, Alabama River at Yellow Jacket Bar, river mile 129.8, 8 March 1967," RDS 4097, Suttkus and Gerald E. Gunning. Paratypes: TU 44028, 7324 specimens taken with holotype; TU 33381, 1613 speci- mens from Alabama, Dallas County, Alabama River at Watts Bar, 3.5 miles upstream from Cahaba, River Mile 204.5, 29 June 1964, RDS 3515, Suttkus and Environmental Biol- ogy Class; TU 35243, 236 specimens from Alabama, Dallas County, Alabama River at old ferry landing across river from Cahaba, 27-28 June 1964, RDS 3508, Suttkus and Environmental Biology Class; TU 35269, 314 specimens from Alabama, Dallas County, Alabama River at Cahaba where Cahaba River enters Alabama River, 28 June 1964, RDS 3513, Suttkus, Glenn H. Clemmer, James D. Archer and John Van Conner; TU 35323, 28 specimens from Alabama, Clarke County, Alabama River along left bank across river from Choctaw Bluff, 2 July 1964, RDS 3519, collected by Suttkus and Environmental Biology Class; TU 40293, 297 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.8, 7 April 1966, RDS 3857, collected by Suttkus and Gunning; TU 40303, 1020 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 7 April 1966, RDS 3858, collected by Suttkus and Gunning; TU 40320, 491 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.4, 7 April 1966, RDS 3859, collected by Suttkus and Gunning; TU 40335, 243 specimens from Alabama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.3, 7 April 1966, RDS 3860, collected by Suttkus and Gunning; TU 40900, 289 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.8, 28 June 1966, RDS 3918, collected by Suttkus, Gunning and Clemmer; TU 40925, 85 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 28 June 1966, RDS 3919, collected by Suttkus, Gunning and Clemmer; TU 40940, 226 speci- mens from Alabama, Wilcox County, Ala- bama River at Tait Bar, River Mile 122.4, 28 No. Type Specimens of Fishes at T/zLiiie 119 June 1966, RDS 3920, collected by Suttkus, Gunning and Clemmer; TU 40950, 109 specimens from Alabama, Wilcox Ccninty, Alabama River at Wilcox Bar, River Mile 120.3, 29 June 1966, RDS 3921, collected by Suttkus, Gunning and Clemmer; TU 41400, 390 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 4.6 miles east of Yellow Bluff Post Office, 1 July 1966, RDS 3924, collected by Suttkus and Environmental Biol- ogy Class; TU 41608, 284 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.8, 4 Octo- ber 1966, RDS 4011, collected by Suttkus and Gunning; TU 41618, 33 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.7, 4 Octo- ber 1966, RDS 4012, collected by Suttkus and Gunning; TU 41632, 92 specimens from Alabama, Wilcox County, Alabama River at Reeves Bar, River Mile 128.5, 4 October 1966, RDS 4013, collected by Suttkus and Gunning, TU 41639, 187 specimens frt)m Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 4 October 1966, RDS 4014, collected by Suttkus and Gunning; TU 41655, 119 specimens from Alabama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.5, 5 October 1966, RDS 4015, collected by Suttkus and Gunning; TU 41666, 147 specimens from Alabama, Wil- cox County, Alabama River at Ohio Bar, River Mile 111.6, 5 October 1966, RDS 4016, collected by Suttkus and Gunning; TU 41670, 301 specimens from Alabama, Mon- roe County, Alabama River at Stein Island, River Mile 107.5, 5 October 1966, RDS 4017, collected by Suttkus and Gunning; TU 41695, 83 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.8, 5 August 1966, RDS 3945, collected by Suttkus and Gunning; TU 41711, 29 specimens from Alabama, Wilcox County, Alabama River at Evans Lower Bar, River Mile 133, 5 August 1966, RDS 3946, collected by Suttkus and Gunning; TU 41726, 16 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.7, 5 August 1967, RDS 3948, collected by Suttkus and Gunning; TU 41732, 19 specimens from Alabama, Wilcox County, Alabama River at Reeves Bar, River Mile 128.5, 5 August 1966, RDS 3949, collected by Suttkus and Gunning; TU 41745, 63 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 5 August 1966, RDS 3950, col- lected by Suttkus and Gunning; TU 41755, 12 specimens from Alabama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.5, 5 August 1966, RDS 3951, collected by Suttkus and Gunning; TU 41761, 9 speci- mens from Alabama, Wilcox County, Ala- bama River at Ohio Bar, River Mile, 111.6, 5 August 1966, RDS 3952, collected by Suttkus and Gunning; TU 41772, 151 speci- mens from Alabama, Monroe County, Ala- bama River at Stein Island, River Mile 107.5, 5 August 1966, RDS 3953, collected by Suttkus and Gunning; TU 41791, 22 speci- mens from Alabama, Monroe County, Ala- bama River at St. James Bar, River Mile 104, 5 August 1966, RDS 3954, collected by Suttkus and Gunning; TU 41797, 4 speci- mens from Alabama, Monroe County, Ala- bama River, at Bates Bar, River Mile 99, 5 August 1966, RDS 3955, collected by Suttkus and Gunning; TU 41813, 10 speci- mens from Alabama, Monroe County, Ala- bama River at Silver Creek Bar, River Mile 87.6, 6 August 1966, RDS 3957, collected by Suttkus and Gunning; TU 41823, 1 speci- men from Alabama, Monroe County, Ala- bama River at mouth of Limestone Creek, River Mile 80.1, 6 August 1966, RDS 3958, collected by Suttkus and Gunning; TU 42731, 370'' specimens from Alabama, Wil- cox County, Alabama River at Evans Upper Bar, River Mile 135.8, 19 December 1966, RDS 4065, collected by Suttkus and Gun- ning; TU 42737, 5630 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.5, 19 De- cember 1966, RDS 4066, collected by Suttkus and Gunning; TU 42746, 687 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 19 December 1966, RDS 4067, collected by Suttkus and Gunning; TU 42759, 900 speci- mens from Alabama, Wilcox County, Ala- bama River at Wilcox Bar, River Mile 120.5, 19-20 December 1966, RDS 4068, collected by Suttkus and Gunning; TU 44011, 175 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River ■■■ Gi\en incorrecth in Suttku.s and Clemmer, 1968: 10 as 470 specimens. 120 Tulane Studies in Zoology and Botany Vol. 16 Mile 135.8, 8 March 1967, RDS 4096, col- lected by Suttkus and Gunning; TU 44029, 94 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 9 March 1967, RDS 4098, collected by Suttkus and Gunning; TU 44045, 561^ specimens from Alabama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.5, 9 March 1967, RDS 4099, collected by Suttkus and Gunning; TU 46783, 202 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.8, 31 May 1967, RDS 4143, col- lected by Suttkus and Gunning; TU 46796, 395 specimens from Alabama, Wilcox County, Alabama River at bar above Yellow Jacket Bar, 31 May 1967, RDS 4144, col- lected by Suttkus and Gunning; TU 46802, 223 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 31 May 1967, RDS 4145, collected by Suttkus and Gunning; TU 46819, 43 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 31 May 1967, RDS 4146, col- lected by Suttkus and Gunning; TU 46830, 32 specimens from Alabama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.5, 31 May-1 June 1967, RDS 4147, col- lected by Suttkus and Gunning; TU 47346, 64 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.7, 7 August 1967, RDS 4177, col- lected by Suttkus and Gunning; TU 47361, 49 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 7 August 1967, RDS 4178, col- lected by Suttkus and Gunning; TU 47374, 86 specimens from Alabama, Wilcox County, Alabama River at Reeves Bar, River Mile 128.5, 7 August 1967, RDS 4179, collected by Suttkus and Gunning; TU 47387, 168 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 7-8 August 1967, RDS 4180, collected by Suttkus and Gunning; TU 47396, 33 speci- mens from Alabama, Wilcox County, Ala- bama River at Wilcox Bar, River Mile 120.6, 8 August 1967, RDS 4181, collected by Suttkus and Gunning; TU 47408, 475 speci- mens from Alabama, Wilcox County, Ala- bama River at Ohio Bar, River Mile 111.7, ^ Given incorrectly in Suttkus and Clemmer, 1968: 20 as 461 specimens. 8 August 1967, RDS 4182, collected by Suttkus, Clemmer and Gunning; TU 47420, 173 specimens from Alabama, Monroe County, Alabama River at Stein Island, River Mile 107.5, 8 August 1967, RDS 4183, col- lected by Suttkus and Gunning; TU 47435, 229 specimens from Alabama, Monroe County, Alabama River at Bates Bar, River Mile 99.4, RDS 4184, collected by Suttkus and Gunning; TU 47452, 134 specimens from Alabama, Monroe County, Alabama River at Haines Island, River Mile 96, 8 August 1967, RDS 4185, collected by Suttkus and Gunning; TU 47477, 89 speci- mens from Alabama, Wilcox County, Ala- bama River at Evans Lower Bar, River Mile 133, 9 August 1967, RDS 4186, collected by Suttkus and Gunning; TU 47491, 24 speci- mens from Alabama, Monroe County, Ala- bama River at St. James Bar, River Mile 104, 10 August 1967,' RDS 4187, collected by Suttkus and Gunning; TU 47499, 16 speci- mens from Alabama, Monroe County, Ala- bama River at Silver Creek Bar, River Mile 87.7, 17 August 1967, RDS 4188, collected by Suttkus and Clemmer; TU 47515, 306 specimens from Alabama, Wilcox County, Alabama River at Hobbs Bar, River Mile 149.5, 18 August 1967, RDS 4189, collected by Suttkus and Clemmer; TU 47762, 143 specimens from Alabama, Wilcox County, Alabama River at Lower Canton Bar, River Mile 156.7, along left bank, 18 August 1967, RDS 4190, collected by Suttkus and Clem- mer; TU 47781, 9 specimens from Alabama, Wilcox County, Alabama River at Lower Canton Bar, River Mile 156.7, along right bank, 18 August 1967, RDS 4191, collected by Suttkus and Clemmer; TU 47822, 85 specimens from Alabama, Wilcox County, Alabama River at Hurricane Island, River Mile 166.5, 19 August 1967, RDS 4193, col- lected by Suttkus and Clemmer; TU 47838, 75 specimens from Alabama, Wilcox County, Alabama River at St. John Bar, River Mile 164.8, 19 August 1967, RDS 4192, collected by Suttkus and Clemmer; TU 47909, 152 specimens from Alabama, Wilcox County, Alabama River at Evans Upper Bar, River Mile 135.7, 26 September 1967, RDS 4204, collected by Gunning and Armand Kuris, TU 47924, 105 specimens from Alabama, Wilcox County, Alabama River at Yellow Jacket Bar, River Mile 129.8, 26 September No. 3 Type Specimens of Fishes at T/ilane 121 1967, RDS 4203, collected by Suttkus and Clemmer; TU 47936, 39 specimens from Alabama, Wilcox County, Alabama River at Reeves Bar, River Mile 128.5, 26 September 1967, RDS 4204, collected by Suttkus and Clemmer; TU 47968, 230 specimens from Alabama, Wilcox County, Alabama River at Tait Bar, River Mile 122.5, 27 September 1967, RDS 4207, collected by Suttkus and Kuris; TU 47980, 108 specimens from Ala- bama, Wilcox County, Alabama River at Wilcox Bar, River Mile 120.6, 27 September 1967, RDS 4208, collected by Suttkus and Kuris; TU 47994, 65 specimens from Ala- bama, Wilcox County, Alabama River at Ohio Bar, River Mile 111.7, 27 September 1967, RDS 4209, collected by Suttkus and Kuris; TU 48012, 232 specimens from Ala- bama, Monroe County, Alabama River at Stein Island, River Mile 107.5, 27 Septem- ber 1967, RDS 4210, collected by Suttkus and Clemmer. Notropis euryzoniis Suttkus, 1955, Tulane Stud. Zool., 3(5):83-100. Paratypes: TU 10700, 8 specimens from Alabama, Russell County, tributary to Little Uchee Creek, 1.1 miles east of Crawford, Highway 80, 17 September 1955, RDS 2520, collected by Suttkus; TU 10718, 60 speci- mens from Alabama, Russell County, tribu- tary to Uchee Creek, 3.2 miles west of Craw- ford, Highway 80, 17 September 1955, RDS 2521, collected by Suttkus. Notropis hypsilepis Suttkus and Raney, 1955, Tulane Stud. Zool., 2(7) : 159-170. Paratypes: TU 8883, 4 specimens (for- merly part of Cornell University No. 17138 ) Georgia, Fulton County, Vickery Creek at junction with Chattahoochee River at Ros- well, Highway 19, 30 March 1950, ECR 1765 and RDS 1765, collected by Edward C. Raney, Suttkus, C. Richard Robins and Richard Backus. Notropis imeldae Cortes 1966, An. Esc. nac. Cienc. biol., Mex., 15:185-192. Paratypes: TU 55375, 5 specimens from Mexico, Oaxaca, Rio Atoyac at Juchatengo, latitude 16° 20' north, longitude 97' 05' west, 16 April 1965, collected by Imelda Martinez and Lauro Gonzalez. Notropis signipinnis Bailey and Suttkus, 1952, Occ. Pap. Mus. Zool. Univ. Mich., No. 542:1-15. Paratypes: TU 1133, 164 specimens from Mississippi, George County, Maletts Spring Branch, tributary to Rocky Creek, tributary to Escatawpa River, 6.6 miles southeast of Lucedale, 4 June 1951, RDS 2147, collected by Suttkus and Hancock. Notropis uranoscopus Suttkus, 1959, Copeia, 1959(1):7-11. Holotype: TU 15289, from Alabama, Bibb County, Cahaba River, 2.2 miles north of Centerville, 17 March 1957, RDS 2594, col- lected by Suttkus, Rudolph J. and Helen V. Miller, John DeAbate. Paratypes: TU 18551, 18 specimens from same collection as holotype; TU 18569, 2 specimens from Alabama, Bibb County, Cahaba River, 2 miles south of Centerville, 5 July 1954, University of Alabama Number 406, collected by Henry Howell, Bryan brothers, Ralph Chermock, A. F. Hemphill, and Herbert Boschung; TU 18570, 8 speci- mens from Alabama, Bibb County, Cahaba River, 2 miles south of Centerville, 5 July 1954, collected by Jack Mallory; TU 18674, 8 specimens from Alabama, Bibb County, Cahaba River, 8.5 miles north of Centerville, Highway 27 crossing, 8 July 1958, RDS 2738, collected by Suttkus, Myrna Andersson, Susan Salisbury, and John DeAbate; TU 19410, 21 specimens from Alabama, Bibb County, Cahaba River, 8.5 miles north of Centerville, Highway 27 crossing, 8 Septem- ber 1958, RDS 2760, collected by Suttkus and Myrna Andersson. Ictaluridae Ictahirus serracanthus Yerger and Relyea, 1 968, Copeia, 1 968 ( 2 ) : 36 1 -384. Paratypes: TU 22592, 4 specimens from Florida, Gadsden County, Lake Talquin just west of mouth of Little River, 20 October 1959, JMB 78, collected by James M. Barku- loo, et al.; TU 22642, 1 specimen from Florida, Leon County, Ochlockonee River at Jackson Bluff, Highway 20, 14-15 Novem- ber 1959, RDS 2861, collected by Suttkus, Barkuloo and Ernest Grover; TU 37226, 3 specimens from Florida, Wakulla County, Sopchoppy River, 2.4 miles south of Sop- 122 Tidane SUidies in Zoology and Botany Vol. 16 choppy, Highway 319, 27 May 1964, col- lected by Kenneth Relyea. Not /mis dlbater. Taylor, 1969, Bull. U. S. Nat. Mus., (282): 1-315. Paratypes: TU 10180, 1 specimen from Arkansas, Newton County, Big Buffalo River, 15.8 miles west of Jasper, Highway 21, 30 April 1955, RDS 2485, collected by Suttkus, Ann Harper, Wanda Hildebrand and Hinton D. Hoese; TU 12240, 15 specimens from Arkansas, Washington County, West Fork White River 6.1 miles south of Fayetteville, Highway 71, 2 September 1955, RDS 2509, collected by Suttkus; TU 12258, 85 speci- mens from Arkansas, Washington County, White River 8.3 miles east of Springdale, Highway 68, 3 September 1955, RDS 2512, collected by Suttkus; TU 16553, 35 speci- mens from Arkansas, Washington County, White River 8.3 miles east of Springdale, Highway 68, 4 October 1957, collected by Suttkus, Charles F. Cole, John DeAbate, Susan Harvey, Myrna Andersson, and Banga- lore I. Sundararaj; TU 16575, 12 specimens from Arkansas, Washington County, White River 8.3 miles east of Springdale, Highway 68, 5 October 1957, RDS 2667, collected by Suttkus, Cole, DeAbate, Harvey, Andersson, and Sundararaj; TU 22401, 7 specimens from Arkansas, Madison County, War Eagle Creek, 5.4 miles north of Huntsville, High- way 23, 25 October 1959, RDS 2856, col- lected by Suttkus, Andersson, Sundararaj; TU 22504, 37 specimens from Arkansas, Newton County, Buffalo Fork of White River, 15.5 miles west of Jasper, Highway 74, 24 October 1959, RDS 2853, collected by Suttkus, Andersson, Sundararaj; TU 22514, 2 specimens from Arkansas, Newton County, Buffalo Fork of White River 15.5 miles west of Jasper, Highway 74, 25 Octo- ber 1959, RDS 2854, collected by Suttkus, Andersson, and Sundararaj; TU 22709, 9 specimens from Arkansas, Cleburne County, Devils Fork Little Red River between Stark and Edgemont, 23 October 1959, RDS 2852, collected by Suttkus, Andersson, and Sun- dararaj. Noturus elegans Taylor, 1969, Bull. U. S. Nat. Mus., (282): 1-315. Paratypes: TU 19390, 12 specimens from Kentucky, Green County, Green River 3.8 miles east-northeast of Donnansburg, High- way 88, 7 September 1958, RDS 2756, col- lected by Suttkus and Myrna Andersson; TU 19460, 16 specimens from Kentucky, Barren County, tributary to Barren River, 2.6 miles southwest of Lucas, Highway 3 IE, at old mill dam, 7 September 1958, RDS 2757, collected by Suttkus and Andersson. Noturus lachneri, Taylor, 1969, Bull. U. S. Nat. Mus., (282): 1-315. Paratypes: TU 7074, 1 specimen from Arkansas, Garland County, 7.7 miles south- southwest of Owensville, 3 miles southeast of U. S. Highway 70, 7 August 1953, EL 94, collected by Ernest Liner; TU 7077, 2 speci- mens from Arkansas, Garland County, 7.7 miles south-southwest of Owensville, 3 miles southeast of U. S. Highway 70, 7 August 1953, EL 94, collected by Ernest Liner. Noturus munitus Suttkus and Taylor, 1965, Proc. Biol. Soc. Wash., 78:169-178. Holotype: TU 26250, from Mississippi, Marion County, Pearl River, 216 miles east of Sandy Hook, 14 November 1955, col- lected by Suttkus and Kristin T. Nielsen. Paratypes: TU 11311, 195 specimens from same collection as holotype; TU 1844, 23 specimens from Mississippi, Marion County, Pearl River 2.3 miles east of Sandy Hook, 24-25 November 1951, RDS 2207, collected by Suttkus and Paul K. Anderson; TU 3557, 8 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 16 February 1952, RDS 2238, col- lected by Suttkus, Paul K. Anderson, Howard K. Suzuki, Thomas Cain; TU 3627, 73 speci- mens from Mississippi, Marion County, Pearl River, 4 miles northeast of Sandy Hook, 13 January 1952, RDS 2217, collected by Suttkus, Anderson and Walter D. Stone; TU 3951, 11 specimens from Mississippi, Marion County, tributary to Pearl River, 4.8 miles southeast of Cokmibia, Highway 13, 23 May 1952, RDS 2297, collected by Suttkus, Cain and Franklin Sogandares-Bernal; TU 4872, 1 specimen from Mississippi, Marion County, tributary to Pearl River, 4.3 miles south of Foxworth, 19 August 1952, RDS 2173, collected by Suttkus, Hancock and Joe Black; TU 7369, 1 specimen from Louisiana, Washington Parish, Pushepatapa Creek, 0.8 mile south of Varnado, 19 January 1952, RDS 2222, collected by Suttkus and Suzuki; No. 3 Type Specimens of Fishes at T/ilane 123 TU 8886, 4 specimens from Mississippi, Marion County, tributary to Pearl River, 0.5 mile south of Hub or 8.5 miles south of Co- lumbia, 28 October 1950, RDS 2013, col- lected by Suttkus and Clarence Lavett Smith; TU 9709, 3 specimens from Mississippi, Marion County, tributary to Pearl River, 5 miles south of Columbia, 30 June 1954, RDS 2456, collected by Suttkus; TU 10462, 3 specimens from Louisiana, Washington Parish, Pushepatapa Creek, 0.8 mile south of Varnado, 29 June 1954, RDS 2455, collected by Suttkus and Nielsen; TU 11588, 1 speci- men from Louisiana, Washington Parish, Pushepatapa Creek, 8.2 miles north of Boga- lusa, 7 May 1956, RDS 2556, collected by Suttkus and Field Zoology Class; TU 13954, 214 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 5 October 1956, RDS 2565, collected by Suttkus, Rudolph J. Miller and Robert K. Chipman; TU 14930, 67 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 12-13 January 1957, RDS 2576, collected by Suttkus and Miller; TU 14938, 82 specimens from Mississippi, Marion County, Pearl River, 2.6 miles east of Sandy Hook, 6 January 1957, RDS 2574, collected by Suttkus and Miller; TU 15073, 2 specimens from Louisiana, Washington Parish, Pushepatapa Creek, 1 mile south of Varnado or 7 miles north of Bogalusa, 13 January 1957, RDS 2578, col- lected by Suttkus and Miller; TU 15171, 92 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 22-23 January 1957, RDS 2580, collected by Suttkus and Norman C. Negus; TU 15465, 2 specimens from Louisiana, Washington Parish, Pushepatapa Creek, 7 miles north of Bogalusa, Highway 21, 21 April 1957, RDS 2600, collected by Suttkus, Robert H. Gibbs, et al.; TU 16148, 6 specimens from Missis- sippi, Marion County, Ten Mile Creek, trib- utary to Pearl River, 9.7 miles north of Sandy Hook, 7 August 1957, RDS 2641, collected by Suttkus and Miller; TU 17472, 1 speci- men from Louisiana, Washington Parish, Pushepatapa Creek, 0.8 mile south of Var- nado, Highway 21, 20 March 1958, RDS 2692, collected by Suttkus, Myrna Andersson and Susan Harvey; TU 17721, 4 specimens from Mississippi, Simpson County, Strong River, 2 miles west of Pinola, 3 April 1958, RDS 2697, collected by Suttkus, Jayson S. Suttkus and Andersson; TU 17951, 1 speci- men from Mississippi, Marion County, Ten Mile Creek, 10.4 miles south of Columbia, Highway 35, 3 May 1958, RDS 2711, col- lected by Suttkus, Andersson and Sidney Simpson; TU 18739, 2 specimens from Mis- sissippi, Simpson County, Strong River, 2 miles west of Pinola, Highway 28 (formerly Hwy 20), 3 July 1958, RDS 2735, collected by Suttkus, Andersson, and J. S. Suttkus; TU 18963, 60 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 3 November 1957, R|M 1053, collected by R. J. Miller, Helen "v. Miller, Andersson and Donald Boyer; TU 19781, 9 specimens from Mississippi, Simp- son County, Strong River, 2 miles west of Pinola, Highway 20, 8 May 1959, RDS 2774, collected by Suttkus and J. S. Suttkus; TU 19825, 52 specimens from Louisiana, Wash- ington Parish, Bogue Chitto River, 1 mile west of Warnerton, 2 May 1959, RDS 2770, Suttkus, Douglas Landwehr, and L. Redman; TU 19880, 3 specimens from Louisiana, Washington Parish, Bogue Chitto River, 1 mile west of Warnerton, 3 May 1959, RDS 2773, Suttkus and Vertebrate Natural His- tory Class; TU 22824, 38 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 26 October 1959, RDS 2857, collected by Suttkus, Andersson, and Bangalore I. Sundararaj; TU 23127, 1 specimen from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 24 March 1960, RDS 2897, collected by Suttkus and Gerald E. Gunning; TU 23310, 1 specimen from Louisiana, Washington Parish, Bogue Chitto River, 1 mile south of Enon, Highway 437, 3 July I960, RDS 2913, collected by Suttkus and John S. Ramsey; TU 23351, 1 specimen from Louisiana, Washington Parish, Bogue Chitto River, 2 miles west of Warnerton, Highway 438, 3 July I960, RDS 2914, collected by Suttkus and Ramsey; TU 23518, 24 specimens from Mississippi, Copiah County, Copiah Creek, 2.4 miles south of Georgetown, Highway 27, 22 July 1960, RDS 2935, collected by Suttkus and Ramsey; TU 23701, I specimen from Mississippi, Marion County, tributary to Pearl River, 0.5 mile north of Marion County line, 21 July 1960, RDS 2928, col- lected by Suttkus and Ramsey; TU 23869, 124 Tulane Studies in Zoology and Botany Vol. 16 3 specimens from Mississippi, Copiah County, Copiah Creek, 2.1 miles south of George- town, Highway 27, 25 November I960, RDS 2940, collected by Suttkus and Prudence Menzies; TU 26657, 87 specimens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 20 October 1961, JSR 61-95, collected by Ramsey and Jamie Thomerson; TU 26695, 1 specimen from Mississippi, Marion County, Holiday Creek, 0.5 mile south of Goss, Highway 13, 21 October 1961, JSR 61-98, collected by Ram- sey and Thomerson; TU 26759, 11 speci- mens from Mississippi, Lawrence County, Fair River, 4.5 miles north of Monticello, Highway 27, 3 November 1961, JSR 61- 104, collected by Ramsey and Thomerson; TU 26878, 1 specimen from Mississippi, Marion County, Silver Creek at north limits of Morgantown, Highway 587, 9 December 1961, JSR 61-116, collected by Ramsey and Thomerson; TU 27137, 5 specimens from Mississippi, Lawrence County, Pearl River, 2.5 miles south of Oakvale, 29 August 1962, RDS 3166, collected by R. D. and J. S. Suttkus; TU 27172, 18 specimens from Mis- sissippi, Marion County, 1.5 miles southeast of Morgantown, 30 August 1962, RDS 3168, collected by R. D. and J. S. Suttkus; TU 27299, 12 specimens from Mississippi, Law- rence County, Bahala Creek, 6 miles south of Rockport, Highway 27, 28 October 1961, JSR 61-103, collected by Ramsey and Thomerson; TU 28034, 87 specimens from Mississippi, Marion County, Pearl River, 1.5 miles southeast of Foxworth, 1 December 1962, RDS 3187, collected by R. D. and J. S. Suttkus; TU 28272, 47 specimens from Mississippi, Marion County, Pearl River, 1.5 miles southeast of Foxworth, 17 October 1962, RDS 3172, collected by Suttkus and Clyde D. Barbour; TU 28303, 4 specimens from Mississippi, Marion County, Pearl River, 3 miles southeast of Foxworth, 18 October 1962, RDS 3173, collected by Suttkus and Barbour; TU 28316, 13 speci- mens from Mississippi, Marion County, Pearl River above mouth of Ten Mile Creek, 6 miles southeast of Foxworth, 18 October 1962, RDS 3174, collected by Suttkus and Barbour; TU 28345, 10 specimens from Mississippi, Marion County, Pearl River, 4 miles north-northeast of Sandy Hook, 0.5 mile upstream from mouth of Hurricane Creek, 18 October 1962, RDS 3175, collected by Suttkus and Barbour; TU 28373, 7 speci- mens from Mississippi, Marion County, Pearl River, 2 miles northeast of Sandy Hook, 18 October 1962, RDS 3176, collected by Suttkus and Barbour; TU 28391, 23 speci- mens from Mississippi, Marion County, Pearl River, 1.5 miles northeast of Sandy Hook, 19 October 1962, RDS 3177, collected by Suttkus and Barbour; TU 28403, 16 speci- mens from Mississippi, Marion County, Pearl River, 2.3 miles east-southeast of Sandy Hook, 19 October 1962, RDS 3178, collected by Suttkus and Barbour; TU 28429, 24 speci- mens from Louisiana, Washington Parish, Pearl River, 4 miles east of Angie, 19 Octo- ber 1962, RDS 3179, collected by Suttkus and Barbour; TU 28534, 240 specimens from Mississippi, Marion County, Pearl River, 1.5 miles southeast of Foxworth, 19-20 Decem- ber 1962, RDS 3188, collected by Suttkus and William T. Mason; TU 28719, 30 speci- mens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 21 October 1961, JSR 61-97, collected by Ram- sey and Thomerson; TU 28768, 129 speci- mens from Mississippi, Marion County, Pearl River, 2.3 miles east of Sandy Hook, 28 De- cember 1962, RDS 3191, collected by Suttkus and Ramsey; TU 28820, 14 specimens from Mississippi, Simpson County, Strong River, 2 miles west of Pinola, Highway 28, 29 De- cember 1962, RDS 3193, collected by Suttkus and Ramsey; TU 29942, 5 specimens from Louisiana, Washington Parish, Pearl River just below sill at Pools Bluff, 4 miles south of Bogalusa, 30-31 August 1963, collected by Suttkus and Gunning; TU 29962, 1 speci- men from Louisiana, St. Tammany Parish, Pearl River at River Mile 52, 0.2 mile below mouth of McGehee Creek, 10 September 1963, RDS 3326, collected by Suttkus, Ram- sey, and Gunning; TU 29975, 1 specimen from Louisiana, St. Tammany Parish, Pearl River at River Mile 46, 10 September 1963, RDS 3328, collected by Suttkus, Ramsey and Gunning; TU 29989, 1 specimen from Loui- siana, St. Tammany Parish, West Pearl River, 5 miles southeast of Talisheek, 1 1 September 1963, RDS 3332, collected by Suttkus, Ram- sey, and Gunning; TU 30010, 17 specimens from Louisiana, Washington Parish, Pearl River just below sill at Pools Bluff, 4 miles south of Bogalusa, 29 September 1963, RDS No. 3 Type Specimens of Fishes at Tulane 125 3342, collected by Suttkus, Ramsey and Gunning; TU 30025, 2 specimens from Louisiana, St. Tammany Parish, Pearl River, 0.2 mile below mouth of McGehce Creek, 10 October 1963, RDS 3343, collected by Suttkus and Gunning; TU 30040, 2 speci- mens from Louisiana, Washington Parish, Pearl River just below sill at Pools Bluff, 4 miles south of Bogalusa, 25 October 1963, RDS 3353, collected by Suttkus and Gunning; TU 30149, 32 specimens from Mississippi, Simpson County, Strong River at rapids up- stream of Highway 28 bridge, 2 miles west of Pinola, 6 November 1963, RDS 3357, col- lected by Suttkus and Ramsey. Noturus phaeus Taylor, 1969, Bull. U. S. Nat. Mus., (282): 1-315. Paratypes: TU 3037, 4 specimens from Mississippi, Lafayette County, tributary to Yocona River, 5.1 miles east of Yocona or 14.7 miles east of Oxford, 23 May 1952, col- lected by Suttkus, Thomas Cain, Franklin Sogandares-Bernal; TU 3124,'' 4 specimens from Mississippi, Lafayette County, tribu- tary to Yocona River, 8 miles east of Oxford, Highway 6, 24 May 1952; TU 14014, 1 specimen from Mississippi, Lafayette County, Yocona River 7.9 miles southeast of Oxford, Highway 334, 12 July 1956, collected by George H. Penn and Joe B. Black; TU 14024, 1 specimen from Mississippi, Alcorn County, 3 miles north of Biggersville, U. S. Highway 45, 12 July 1956, collected by Penn and Black; TU 14036, 2 specimens from Mississippi, Benton County, 0.3 mile west of Michigan City, Mississippi, Highway 7, 11 July 1956, collected by Penn and Black. Noturus stigrnosus Taylor, 1969, Bull. U. S. Nat. Mus., (282): 1-315. Paratypes: TU 19217, 4 specimens from Michigan, Washtenaw County, Huron River at North Territorial Road, 4 miles northwest of Dexter, 31 August 1958, RDS 2750, col- lected by Suttkus, Carter Gilbert, and Wil- liam Davis; TU 19246, 6 specimens from Michigan, Washtenaw County, Huron River at North Territorial Road, 4 miles northwest of Dexter, 3 September 1958, RDS 2752, col- lected by Suttkus, Myrna Andersson, and Thomas Poulson. " Given incorrectly in Taylor, 1960: iOi a.s TU 3134. Prietella phreatophila Carranza, 1954, Cien- cia, Mex., 14 ( 7-8 ): 129-136. Paratypes: TU 10810, 4 specimens (for- merly in the series retained by Carranza at time of publication, subsequently sent to Tulane as a gift; one specimen removed, skele- tonized and recataloged as Osteology Collec- tion No. 178) from Mexico, Coahuila, "El Potrero" de Doua Mariana, Municipio de Muzquiz, at the base of Sierra de Santa Rosa, 29 August 1954, collected by Jorge Carranza and C. Bolivar. Cyprinodontidae Cualac tessellatus Miller, 1956, Occ. Pap. Mus. Zool. Univ. Mich., 581 : 1-17. Paratypes: TU 6557, 32 specimens from Mexico, San Luis Potosi, La Media Luna, about 7 miles south-southwest of settlement of Rio Verde, collected by Richard T. Gregg. Caproidae Antigonia comhatia Berry and Rathjen, 1959, Quart. Journ. Fla. Acad. Sci. (1958; 21(3):255-258. Paratypes: TU 18373, 2 specimens from Atlantic Ocean, latitude 07 " 31' north, longi- tude 54^ 16' west, off Surinam, 125 fathoms, 8 November 1958, Oregon Sta. No. 2013, collected by M/V Oregon. Atherinidae Chirostoma riojai Solorzano and Lopez, 1965, Ciencia, Mex. 24 ( 3-4 ) : 1 45-150. Paratypes: TU 40173, 2 specimens from Mexico, Mexico, Laguna de Santiago Tilapa, 31 October 1963, LB 1820, collected by A. Solorzano, R. Cruz, R. Aguilar and F. Car- mona. Poblana jerdebueni Solorzano and Lopez, 1965, Ciencia, Mex., 23 ( 5 ) :209-213. Paratypes: TU 38631, 3 specimens from Mexico, Puebia, Laguna de Almoloya o Chignahuapan, 22 May 1959, L B. 951, collected by A. Solorzano and T. Obregon. Serranidae Pikea mexicanus Schultz, 1958, Proc. U. S. Nat. Mus., 1 08 ( 3405 ) : 32 1-329. Paratypes : TU 1 1 004, 6 specimens from Gulf of Mexico, latitude 27^ 10' north. 126 Tulane Studies in Zoology and Botany Vol. 16 longitude 96° 20' west, 150 fathoms, 5 June 1954, Oregon Sta. 1094, collected by M V Oregon; TU 11008, 1 specimen from Gulf of Mexico, latitude 27° 10' north, longitude 96° 20' west, 150 fathoms, 5 June 1954, Oregon Sta. 1094, collected by M V Oregon; TU 12972, 11 specimens from Gulf of Mexico, latitude 28° 47' north, longitude 85° 19' west, 64 fathoms, 40 foot flat trawl, 7 March 1953, Oregon Sta. 895, collected by M/V Oregon; TU 12985, 1 specimen from Gulf of Mexico, latitude 29° 19' north, longitude 86° 04' west, 82 fathoms, 40 foot trawl, 31 October 1953, Oregon Sta. 864, col- lected by M/V Oregon; TU 14739, 1 speci- men from Gulf of Mexico, latitude 27° 40' north, longitude 95° 45' west, 100 fathoms, 80 foot balloon trawl, 7 May 1956, Oregon Sta. 1510, collected by M/V Oregon. Centrarchidae Ambloplites anommus Viosca 1936, Copeia, 1936(1 ) :37-45 (currently considered as subspecies, Ambloplites rupestris ariom- nius ) . Paratypes: TU 2856, 14 specimens from Louisiana, St. Tammany Parish, Little Bogue Falaya "Falia" Creek, 3 miles north of Coving- ton, 19 May 1935, collected by Percy Viosca and George H. Penn ( formerly in the private collection of the late Percy Viosca; given to Tulane about 1955; specimens are green be- cause the original label was a shipping tag with a copper eyelet and the specimens were kept in formalin by Viosca). Sparidae Calamus nodosus Randall and Caldwell, 1966, Bull. Los Angeles County Mus. Nat. Hist. Sci., (2): 1-47. Paratype: TU yilG'b, 1 specimen from Atlantic Ocean off Georgia Coast, latitude 33° 42' north, longitude 77° 11' west, 12-20 fathoms, 9 December 1959, Silver Bay Sta. 1507, collected by M/V Silver Bay. Percidae Etheostoma collettei Birdsong and Knapp, 1969, Tulane Stud. Zool. and Bot., 15(3): 106-112. Paratypes: TU 52876, 5 specimens from Louisiana, Union County, Meridian Creek, 1 mile east of Conway, sec. 17, T22N, RIE, 21 May 1956, collected by William R. Taylor and L. W. Lowe. Etheostoma ditrema Ramsey and Suttkus, 1965, Tulane Stud. Zool., 12(3) :65-77. Holotype: TU 35703, from Georgia, Chat- tooga County, a spring flowing into a tribu- tary to Mills Creek, tributary to Chattooga River, 4.3 airline miles due west of Lyerly, 18 July 1962, RDS 3134, collected by Suttkus and William T. Mason. Paratypes: TU 29153, 21 specimens from same collection as holotype; TU 26086, 8 specimens from Georgia, Chattooga County, spring tributary to Mills Creek, tributary to Chattooga River, 5.3 miles west of Lyerly, 19 April 1962, RDS 3052, collected by Suttkus, John S. Ramsey, Jamie Thomerson, and Clyde D. Barbour; TU 27566, 9 specimens from spring tributary to Mills Creek, tribu- tary to Chattooga River, 5.3 miles west of Lyerly, 30 May 1962, JSR 62-57, collected by John S. Ramsey; TU 32981, 43 specimens from Georgia, Chattooga County, spring trib- utary to Mills Creek, tributary to Chattooga River, 5.3 miles west of Lyerly, 23 June 1964, RDS 3499, collected by Suttkus and Environ- mental Biology Class. Etheostoma m,icrolepidum Raney and Zorach, 1967, Am. Midi. Nat., 77( 1 ) :93-103. Paratypes: TU 19479, 19 specimens from Tennessee, Rutherford County, East Fork of Stones River, 6 miles north of Murfreesboro, Highway 231, 7 September 1958, RDS 2758, collected by Suttkus and Myrna Andersson; TU 32983, 71 specimens from Tennessee, Cheatham County, Harpeth River, 8 miles east of White Bluff, Highway 70, 15 July 1964, JSR 64-60, collected by John S. Ram- sey and Dan Walton. Etheostoma moorei Raney and Suttkus, 1964, Copeia, 1964(1):130-139. Paratypes: TU 22697, 41 specimens from Arkansas, Cleburne County, Devils Fork, Little Red River between Stark and Edge- mont, Highway 16, 23 October 1959, RDS 2852, collected by Suttkus, Myrna Andersson, and Bangalore I. Sundararaj; TU 26227, 22 specimens from Arkansas, Van Buren County, Middle Fork, Little Red River at Shirley, 17 August 1962, RDS 3157, collected by Suttkus and William T. Mason; TU 26249, 21 speci- mens from Arkansas, Van Buren County, No. 3 Type Specimens of Fishes at Tidane 111 South Fork, Little Red River, 9 miles south- west of Clinton, Highway 95, 17 August 1%2, RDS 3158, collected by Suttkus and Mason. Etheostoma niichale Howell and Caldwell, 1965, Tulane Stud. Zool., 12(4) : 101-108. Paratypes: TU 34591, 20 specimens from Alabama, Jefferson County, Glen Spring at Bessemer, NE ¥4, Sec. 17, T 19S, R4W, along county Highway 20, 9 September 1964, RDS 3582, collected by Suttkus, John S. Ramsey, and Francis L. Rose. Poecilichtbys wbipplii radiosus Hubbs and Black, 1941, Occ. Papers Mus. Zool. Univ. Mich., 429:1-27. (Now known as Etheo- stoma radiosum ) . Paratypes: TU 52871, 1 specimen from Arkansas, Garland County, North Fork, Oua- chita River, 2.5 miles west of Buckville, 1 June 1939, collected by T. Holder; TU 52872, 5 specimens from Arkansas, Garland County, Cataract Branch, 1% miles north of Cedar Glade, 2 May 1939, collected by T. Holder; TU 52873, 2 specimens from Arkansas, Garland County, Big Branch, 2^/4 miles west of Buckville, I6 May 1939, col- lected by T. Holder. These eight paratypes apparently had been retained at the Univer- sity of Arkansas by John D. Black subsequent to the description. During 1967, a large part of the University of Arkansas' fish collection was given to Tulane University and thus the transfer of the type material. Etheostoma rnbriim Raney and Suttkus, 1966, Tulane Stud. Zool., 13(3 ) : 95-102. Paratypes: TU 30171, 47 specimens from Mississippi, Copiah County, Bayou Pierre, 8.6 miles southwest of Utica, Highway 18, 7 December 1963, JSR 63-58, collected by John S. Ramsey and Michael D. Dahlberg; TU 31341, 2 specimens from Mississippi, Hinds County, White Oak Creek, tributary to Bayou Pierre, 2.7 miles south of Utica, Highway 18, 1 February 1964, RDS 3378, collected by Edward C. Raney and Suttkus; TU 37305, 4 specimens from Mississippi, Copiah County, Bayou Pierre, 8.6 miles south- west of Utica, Highway 18, 18 March 1965, RDS 3643, collected by Suttkus and Larry Ogren; TU 37451, 53 specimens from Mis- sissippi, Copiah County, Bayou Pierre, 8.6 miles southwest of Utica, Highway 18, 28 April 1965, RDS 3661, collected by Suttkus and Francis L. Rose; TU 40368, 10 speci- mens from Mississippi, Copiah County, Bayou Pierre, 8.6 miles southwest of Utica, Highway 18, 2 April 1966, RDS 3855, col- lected by Suttkus and Glenn H. Clemmer. Percina aurolineata Suttkus and Ramsey, 1967, Tulane Stud. Zool., 13(4) : 129-145. Holotype: TU 39573, from Georgia, Gil- mer County, Coosawattee River about 200 yards below the mouth of a small spring tributary, 4.2 miles southwest of the center of Ellijay, 18 June 1966, RDS 3910, col- lected by Suttkus and Environmental Biology Class. Paratypes: TU 41049, 3 specimens from same collection as holotype; TU 38320, 5 specimens from Georgia, Gilmer County, Cartecay River, tributary to Coosawattee River, 2.9 miles southeast of Ellijay, along Highway 52, 18 June 1965, RDS 3695, col- lected by Suttkus and Environmental Biology Class; TU 38507, 15 specimens from Georgia, Gilmer County, Coosawattee River, 4.2 miles southwest of Ellijay, 22 June 1965, RDS 3707, collected by Suttkus and Environmen- tal Biology Class; TU 40714, 4 specimens from Georgia, Gilmer County, tributary to Cartecay and Cartecay River, 2.9 miles south- east of Ellijay, along Highway 52, 15 April 1966, RDS 3881, collected by Suttkus and Glenn H. Clemmer; TU 41053, 6 specimens from Georgia, Gilmer County, Cartecay River, 3.1 miles southeast of Ellijay, along Highway 52, 17 June 1966, RDS 3909, col- lected by Suttkus, Clemmer, and Environ- mental Biology Class. Percina lenticula Richards and Knapp, 1964, Copeia, 1964, (4):69()-701. Paratypes: TU 15291, 7 specimens from Alabama, Bibb County, Cahaba River, 2.2 miles north of Centerville, just off Highway 5, 17 May 1957, RDS 2594, collected by Suttkus, Rudolph J. and Helen V. Miller, and John DeAbate. Clinidae Starksia y din eat a Gilbert, 1965, Notulae Naturae, No. 379:1-6. Paratypes: TU 36401, 2 specimens from Caribbean Sea, Grand Cayman Island, "Iron- shore" formation in front of Seaview Lodge, 128 Tulane Studies in Zoology and Botany Vol. 16 south side of Georgetown, depth 2-12 feet, collected by Carter R. Gilbert and John C. Tyler. Microdesmidae Microdesmus carri Gilbert, 1966, Copeia, 1966, {2):325-332. Paratypes: TU 36400, 2 specimens from Atlantic Ocean, Costa Rica, Limon Province, east side of Tortiiguero lagoon, near mouth, 7 August 1964, collected by D. P. Kelso. Microdesmus suttkiisi Gilbert, 1966, Copeia, 1966, (2): 325-332. Holotype: TU 25110, from Costa Rica, Puntarenas Province, boca de Rio Baru at Dominical, 28 January 1961, RDS 2961, col- lected by Suttkus, Alfred Smalley, et al. Paratypes: TU 25130, 1 specimen from Costa Rica, Puntarenas Province, Rio Baru, 1 kilometer north of Dominical, 28 January 1961, RDS 2962, collected by Suttkus, Smalley, et al.; TU 25262, 1 specimen from Costa Rica, Puntarenas Province, at La Barca, 27 February I960, collected by Salvador Jimenez-C. Draconettidae Draconetta oregona Briggs and Berry, 1959, Copeia, 1959, (2): 123-133. Paratypes: TU 18831 (not 188831 as given in description), 3 specimens from Atlantic Ocean, latitude 02° 04' north, longitude 47° 00' west, 125 fathoms, 17 November 1957, Oregon Sta. 2080, collected by M V Oregon. Gobiidae Gobiosoma evelynae Bohlke and Robins, 1968, Proc. Acad. Nat. Sci. Philadelphia, 120(3):45-174. Paratypes: TU 48782, 2 specimens from Atlantic Ocean, Virgin Islands, Anguilla Island, Dowlings Shoal, west of Crocus Bay off north end of Sand Island, 17 July 1965, collected by James C. Tyler and William N. Eschmeyer. Gobiosoma genie Bohlke and Robins, 1968, Proc. Acad. Nat. Sci. Philadelphia, 120 (3):45-174. Paratypes: TU 48781, 1 specimen from Atlantic Ocean, Bahama Islands, Hog Island, north shore Cstation 122), 15 March 1953, collected by Charles C. G. Chaplin. Triglidae Bellator ribeiroi Miller, 1965, Quart. Jour. Florida Acad. Sci., 28(3 ) :259-266. Paratypes: TU 37821, 2 specimens from Atlantic Ocean off Colombia, latitude 08° 50.5' north, longitude 76° 53.5' west, 37-40 fathoms, 27 May 1964, Oregon Sta. 4899, collected by M V Oregon. Peristedion greyae Miller, 1967, Bull. Mar. Sci., 17 (U: 16-41. Paratypes: TU 33302, 2 specimens from Atlantic Ocean off North Carolina, latitude 34° 15' north, longitude 75° 54' west, 190- 200 fathoms, 9 June 1962, Silver Bay Sta. 41 59, collected by M V Silver Bay. Cottidae Coitus pygmaeus Williams, 1968, Copeia, 1968, (2):334-342. Paratypes: TU 34396, 47 specimens from Alabama, Calhoun County, Coldwater Creek, just below Coldwater Springs at town of Coldwater, 31 August 1964, RDS 3571, col- lected by Suttkus and John S. Ramsey. Tetraodontidae Sphoeroides parvus Shipp and Yerger, 1969. Proc. Biol. Soc. Wash., 82:477-488. Paratypes: TU 9381, 1 specimen from Louisiana, St. Tammany Parish, Lake Pont- chartrain, 2 miles west of South Drawbridge of L and N railroad, latitude 30" 10' north, longitude 89° 55' west, 5 November 1954, M/V Anna Inez Station 547, collected by Suttkus and party; TU 19038, 2 specimens from Louisiana, Cameron Parish, at jetties along west bank of Calcasieu River at Cameron, 28 April 1957, RFS 7, collected by Rita F. Smith and Michael Guidry; TU 22573, 14 specimens from Louisiana, Jeffer- son Parish, Gulf of Mexico, 100 yards off Grand Terre Island near Grand Isle, 12 De- cember 1959, collected by Douglas Landwehr and Richard D. Lumsden. Batrachoididae Porichthys bathoiketes Gilbert, 1968, Bull. Mar. Sci., 18( 3) :671-730. Paratypes: TU 18572, 9 specimens from Atlantic Ocean, latitude 11° 27' north, longi- tude 83 ' 11' west, 135 fathoms, 9 September No. 3 Type Specimens of Fishes at Tulane 129 1957, Oregon Sta. 1902, collected by M/V Oregon; TU 18573, 1 specimen from Atlan- tic Ocean, latitude 16° 07' north, longitude 81° 05' west, 130 fathoms, 24 August 1957, Oregon Sta. 1891, collected by M V Oregon; TU 18574, 3 specimens from Atlantic Ocean, latitude 16° 38' north, longitude 81° 39' west, 150 fathoms, 22 August 1957, Oregon Sta. 1879, collected by M V Oregon; TU 19928, 2 specimens from Atlantic Ocean, latitude 16° 39' north, longitude 81° 43' west, 125 fathoms, 22 August 1957, Oregon Sta. 1878, collected by M V Oregon. Summary A brief history of the Tulane University fish collection and a list of type specimens are given. The modern collection was started in 1950 and has grown to nearly two million catalogued specimens in two decades. The following seven holotypes are housed in the Tulane University collection: Algansea mon- ticola, Notropis edwardraneyi, Notropis i/ranoscopas, Noturus munitiis, Etheostoma ditrema, Percina aurolineata, Microdesnius suttkiisi. There are 30,190 paratypes of 47 different species. Complete Citations of Original Descriptions HiLDEBRAND, Samuel F. 1948. A review of the American menhaden, genus Bicvooiiia, with a description of a new species. Smithson. Misc. Coll., 107 (18): 1-39. GiBBs, Robert H., Jr. 1960. Alcpimurus brevirostris, a new species of lancet fish from the Western north Atlantic. Breviora (123): 1-14. Barbour, Clyde D., and Salvador Coxtrebas. 1968. Al^anaca monticola, a new cyprinid fish from the Pacific slope of central Mexico. Proc. Biol. Soc. Wash., 81:101-108. Lachner, Ernest A., and Robert E. Jenkixs. 1967. Systematics, distribution and evolution of the chuli genus Nocomis (Cyprinidae) in the southwestern Ohio River basin, with the description of a new species. Copeia, 1967 (3):557-580. SuTTKUs, Royal D., and Edward C. Raxey. 1955. Notropis asperifrons, a new cyprinid fish from the Mobile Bay drainage of Ala- bama and Georgia, with studies of related species. Tulane Stud. Zool., 3(l):l-33. SuTTKUs, Royal D., and Edward C. Raney. 1955. Notropis baileyi, a new cyprinid fish from the Pascagoula and Mobile Bay drain- ages of Mississippi and Alabama. Tulane Stud. Zool., 2(5):69-86. SuTTKUs, Royal D., and Glexn H. Clemmer. 1968. Notropis edwardraneyi, a new cyprinid fish from the Alabama and Tombigbee river systems and a discussion of related species. Tulane Stud. Zool. and Bot., 15( I) : 18-39. SuTTKUs, Royal D. 1955. Notropis euryzontis, a new cyprinid fish from the Chattahoochee River system ot Cieorgia and Alal)aina. 'iulane Stud. Zool., 3(5) :83-100. SuTTKus, Royal D., and Edward C. Raxey. 1955. Notropis hypsilcpis, a new cyprinid fish from the .Xpalachicola River system of Georgia and Alabama. Tulane Stud. Zool., 2(7): 159-170. Cortes, M. T. 1966. Cousideraciones sobre el genero Notropis- y descripcion de ima especie nueva, procedente del rio Atoyac en Jucha- tengo, Oax., Mexico. An. Esc. nac. Cienc. bioL, Max., 15:185-192. Bailey, Reeve M., and Royal D. Suttkus. 1952. Notropis sinui))innis, a new cyprinid fish from southeastern United States. Occ. Pap. Mus. Zool. Univ. Mich., No. 542:1-15. Suttkus, Royal D. 1959. Notropis urano- scopus, a new cyprinid fish from the Alabama River system. Copeia, 1959 (1):7-11. Yergek, Ralph, and Kenneth Relyea. 1968. The flat-headed bulllieads (Pisces: Ictaluridae) of the southeastern United States, and a new species of Ictahints from the Gulf Coast. Copeia, 1968 (2):361-384. Taylor, WiLLi.-\>t Ralph. 1969. A revision of the catfish genus Notiinis Rafinesque with an analvsis of higher groups in the Ictaluridae. Bull. Nat. Mus., No. 282:1-315. Suttkus, Royal D., and William Ralph Tay- lor. 1965. Notunts luunittis. a new species of madtom, family Ictaluridae, from south- em United States. Proc. Biol. Soc. Wash., 78:169-178. Carraxza, Jorge. 1954. Descripcion del primer bagre anoftalmo y depigmentado en- contrado en aguas mexicanus. Ciencia, 14 (7-8): 129-136. Miller, Robert Rush. 1956. A new genus and species of cyprinodontid fish from San Luis Potosi, Mexico, with remarks on the sub- familv Cvprinodontinae. Occ. Pap. Mus. Zool. Univ.' Mich., 581:1-17. Berry, Frederick H., and \\ ahhkn F. Rath j en. 1958. A new species of the hoarfisli genus Antigonia from the western Atlantic. Quart. Journ. Fla. Acad. Sci., 21(3) :255-258. SoLORZANO, A., and Y. Lopez. 1965. Nueva especie de Chirostonia capturada en la La- guna de Victoria o de Santiago Tilapa, Estado de Mexico (Pise, Atherin.). Ciencia, Max., 24(3-4): 145-150. SoLORZAXO, A., and Y. Lopi/. 1965. Nue\a especie de Pohhiiia capturada en la Laguna de Almolova o Chignahuapan, Estado de Puebla (Mexico). Ciencia, Mex., 23(5): 209- 213. ScHULTZ, Leoxard P. 1958. Three new .ser- ranid fishes, genus Pikea. from tlie western AUantic. Proc. U. S. Nat. Nh.s., 108 (3405): 321-329. VioscA, Percy, Jr. 1936. A new rock bass from Louisiana and Mississippi. Copeia, 1936 (l):37-45. ft I Volume 16, Number 4 September 23, 1970 xviUo. COi^..-. _.., LIBRARY OCT 71970 HARVARD UNlVERSiTYi A SYSTEMATIC REVIEW OF UROMACER CATESBYI SCHLEGEL (SERPENTES, COLUBRIDAE) ALBERT SCHWARTZ Department of Biology, Miami-Dade Junior College, Mj'ami, Florida 33167 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, title pages and tables of contents are distributed to institutions receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in chapters I and II (only) of the Style Manual for Biological Journals, 2nd ed., published in 1964 by the American Institute of Biological Sciences, Washington, D. C. Manuscripts should be submitted on good paper, as original typewritten copy, double- spaced, and carefully corrected. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. 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Price per number (flat rate) : $1.50 Price per volume (flat rate) : $5.00 Meade Natural History Library Tulane University Riverside Research Laboratories Route 1, Box 46-B Belle Chasse, Louisiana 70037 Gerald E. Gunning, Editor Alfred E. Smalley, Associate Editor Arthur L. Welden Associate Editor Department of Biology Tulane University New Orleans, Louisiana 70118, U.S.A. Vickie L. Haddenhorst Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY MUS. CO MP. ZOOL. LIBRARY OCT 71970 Volume 16, Number 4 September Fi^,RM)^D — UNIVERSITY. A SYSTEMATIC REVIEW OF UROMACER CATESBYI SCHLEGEL (SERPENTES, COLUBRIDAE) ALBERT SCHWARTZ Department of Biology, Miunu-Dade Junior College, Miami, Florida 33167 Abstract Vromacer eateshtji occurs tlirouuhout Hispaniola and on seven satellite islands. The species is iDolytypic and the followinti subspecies are recognized: U. c. cateshiji (Tiburon Peninsula, ILiiti, west of Mo- mance), L^ c. eereolineaius (Isles Petite and Grande Cayemite, Haiti), U. c. frondi- color (lie de la Conave, Haiti), U. c. hari- olatus ( Haiti, north of the Cul de Sac Plain), L". c. inchausteguii (Isla Saona, Republica Doniinicana ), U. c. insulaerac- carum ( Ile-a-Vache, Haiti), V. c. pampi- neus ( Republica Doniinicana, north of the Valle de Neiba), and V. e. scandax (lie de la Tortue, Haiti). Populations inhabiting the Cul de Sac- Valle de Neilia and the Pe- ninsula de Barahona are assigned intergrade status between cateshiji, hariolatits, and pampineus. The subspecies have differen- tiated in pattern details and in \entral and subcaudal scale counts. Prototype cateshiji presumably divided early into north and south islands populations, the latter with a bold longitudinal line on the lower scale rows. Further differentiation proceeded from these bases; satellite forms are as- sociated with the subspecies on the adja- cent mainland. The colubrid snake genus Uromacer Du- meril and Bibron is one of four endemic colubrid genera on the Antillean island of Hispaniola. Cochran (1941:329 et seq.) recognized six species in the genus: catesbyi Schlegel, 1837; scandax Dunn, 1920; jren- atus Giinther, 1865; wetmorei Cochran, 1931; dorsalis Dunn, 1920; and oxyrhynchus Dumerii and Bibron, 1854. These six species are separable into two groups: one group ( catesbyi, scandax ) has the head only slightly elongate with but little modification in the shape of the preorbital scutellation, whereas the other (the remaining four taxa ) has the head very much attenuate, in the fashion of the better known New World mainland genus Oxybelis. Of the six forms recognized by Cochran, three {scandax. icetyjiorei. dor- salis) were considered to be endemic to His- paniolan satellite islands ( He de la Tortue, Isla Beata, He de la Gonave, respectively). All Uromacer are either green, gray, tan, or some combination of these three basic hues. The long-snouted forms are "vine snakes" in that they are adept climbers and sleep above ground at the tips of branches, whereas the short-snouted forms are apparently somewhat more terrestrial in their activity (but see comments on diet and sites of for- aging by Horn, 1 969 ) . Evidence from field studies indicates that there are fewer than six species of Uromacer. At the time of Cochran's work, many areas of Hispaniola were unrepresented by collec- tions, and these hiatuses in the distribution of members of the genus may have been re- sponsible for Cochran's interpretation of the distinctness of the various taxa. At most, there appear to be three species ( Horn, 1969, recognized four); it is the purpose of this paper to discuss the geographic vari- ation in one of these, U. catesbyi, to which species I assign both of the short-snouted taxa: catesbyi and scandax. I have examined 265 U. catesbyi ( includ- ing U. scandax). Considering the amount of herpetological collecting in Hispaniola in recent years, this is a remarkably small niunber of snakes. There are other specimens Editorial Committee for this Paper: Dr. Edmond V. Malnate, The Academy of Natural Sciences, Nineteenth and the Parkway, Philadelphia, Pennsylvania 19103 Dr. Larry David Wilson, Department of Biology, University of Southwestern Louisiana, Lafayette, Louisiana 70501 131 132 T/ilane Studies in Zoology and Botany Vol. 16 in collections, notably in the United States Coy, Jr., Richard G. Zweifel and George W. National Museum, which I have not studied, Foley, Ernest E. Williams, and James A. since they in general duplicate material ex- Peters, respectively, all of whom I wish to amined in other collections. Nevertheless, thank for their cooperation. A single snake U. catesbyi seems a less abundant (or pos- in the collection of Lewis D. Ober (EDO) sibly less easily secured because of its cryptic has also been available to me. Holotypes and coloration) species than other members of paratypes of new subspecies have been des- the genus. ignated or deposited in the above collections. Collections made in 1963 by myself and I have counted ventral scales in the man- party in Haiti easily demonstrated that U. ner proposed by Dowling (1951). Paired catesbyi from the Tiburon Peninsula west of subcaudal scales also have been tabulated, Port-au-Prince differed markedly in life from but, as in many long-tailed snakes, a large snakes from north of the Cul de Sac Plain. number of V . catesbyi have incomplete tails. Additional specimens from both Haiti and Since the tail is extremely attenuate termi- the Republica Dominicana taken between nally, even the loss of as little as a centimeter 1964 and 1968 nor only confirmed these from the end of the tail may profoundly affect originally observed differences but also dem- the subcaudal count. Accordingly, I have onstrated the existence of other distinct counted the subcaudals of only those snakes populations on Hispaniola and its satellite that still retain the terminal spine. Supra- islands. Although there are obvious differ- and infralabials, loreals, pre- and postoculars, ences in scutellation between these popula- and temporals also were counted. Number tions, the most prominent traits are pig- of dorsal scale rows behind the head, at mid- mental. Data on the coloration of living body, and before the vent were counted, and individuals are invaluable in a study of the each snake was measured ( both snout-vent variation in U. catesbyi. length and tail length ) in millimeters. Specimens collected by myself and various Uro??iacer catesbyi (sensu lata) is widely parties are in the Albert Schwartz Field Series distributed on the Hispaniolan mainland (ASFS). Collections from the Republica and has been taken also on He de la Tortue, Dominicana were made in 1968 under Na- He de la Gonave, He Petite and He Grande tional Science Foundation Grant GB-7977. Cayemite, He-a-Vache, and Isla Saona. There I wish to acknowledge the field assistance are no specimens or records of the species of Patricia H. Adams, Robert K. Bobilin, from Isla Beata, where one of the long- Sixto J. Inchaustegui, Ronald F. Klinikowski, snouted forms of Uromacer occurs, but almost David C. Leber, John K. Lewis, Dennis R. certainly U. catesbyi will be taken on that Paulson, James A. Rodgers, Jr., and Richard island. Although there are no specimens Thomas, all of whom aided immeasurably in from Isla Catalina off the southern Domin- collecting these snakes in Hispaniola. Mr. ican coast, in August 1968 I examined a Thomas deserves special mention for col- much decayed U. catesbyi that had been lecting on He Grande Cayemite for me, and killed by members of the Catalina naval gar- Mr. Klinikowski and Drs. Leber and Paulson rison a few days previously; thus, the snake visited Ile-a-Vache on my behalf. The avail- does occur on that islet. ability of U. catesbyi from Isla Saona is due Uromacer catesbyi is an inhabitant of primarily to the cooperation of Sr. Inchau- wooded situations, but the woods need not tegui, whose assistance in securing trans- be extremely mesic nor high. Presence of portation to that little-known island cannot shade seems to be a predominant factor, and be minimized. In addition to specimens in such lowland situations as lime and cacao the ASFS, I have studied snakes in the col- groves or scrub and cultivated gardens offer lections of the Carnegie Museum ( CM ) , sufficient retreats and shelter for these the American Museum of Natural History snakes. Most individuals secured by us (AMNH), the Museum of Comparative were found on the ground or in low bushes. Zoology at Harvard University ( MCZ ) , A snake was taken at Centro de Boya in and the United States National Museum the Republica Dominicana, however, as it (USNM). These snakes were lent to me crawled across the surface of a termitarium by Neil D. Richmond and Clarence J. Mc- at a height of five meters in a tree adjacent No. 4 Uromacer catesbyi 133 to a road in a patch of broadleaf woods; one catesbyi. On Torttic, C. Rhea Warren also of a pair of snakes ( which were apparently spent two days, and, despite promises of re- fighting ) was secured from a banana tree muneration for green snakes, the natives at Peneau in Haiti. Perhaps the most inter- brought no U. catesbyi and only a very few esting site for collection of U. catesbyi was U. oxyrhynchus. Coupled with the fact that a stand of deciduous woods that borders a only three Tortue U. catesbyi are present in road near Palo Verde in the Republica Do- collections, the experience suggests that on minicana. The woods were separated from Tortue the species is quite uncommon. On the road by a barbedwire fence and were Isla Saona, natives brought us both U. cates- surrounded by land planted in sugar cane. byi and U. oxyrhynchus, more of the latter Five JJ. catesbyi were taken as they traveled species being represented. Judged only by along the barbedwire. When alarmed, the my experience, on He de la Gonave V. dor- snakes glided away rapidly, using the tops salis outnumbers U. catesbyi, but on Ile-a- of low shrubs adjacent to the fence as an Vache U. catesbyi appears to outnumber U. avenue of escape into the woods beyond, frenat/is. Reasons for the abundance or Despite our presence, persistent patrolling rarity of U. catesbyi ( or its relative abun- of the length of fence continued to reveal dance to one of the long-snouted forms) on more snakes, and it seemed certain that even these satellite islands are unknown, though alarmed the snakes returned rather All populations of V . catesbyi are some quickly to lie upon the shaded barbedwire. shade of green both dorsally and ventrally, Anoles were extremely abundant in the with the ventral color lighter ( more yellow woods and on the fence, and I am confident or paler green ) than the dorsal color. Those that the snakes, lying extended on the wire populations associated with the Tiburon Pe- along which the anoles scampered, found ninsula in Haiti (including snakes from Ile-a- this method a simple one of securing food. Vache and the Cayemites ) have a prominent I have not examined stomach contents of U. lateral line in life, usually associated with catesbyi but a Leiocephabis m. melanochloriis scale rows one to three, which is pale blue, was regurgitated by a snake taken on Ile-a- pale greenish, or white. The dorsal scales Vache. Since this lizard regularly climbs usually are very finely edged with black, but (Schwartz, 1966:48), the fact that it serves in the Isla Saona population, the dark scale as a food item for U. catesbyi does not neces- edging is very bold and gives a distinctly sarily indicate terrestrial foraging on the squamate appearance to the dorsum. Ad- part of the snake. Mertens (1939:78) re- ditionally, two populations have dark gray- ported a species of Anolis and Hyla domini- green markings on the dorsal head scales. All censis as food items for this species. Horn these pattern elements are important in dis- (1969:8-9) listed Hyla pulchrilineata, H. tinguishing the constituent subspecies. In dominicensis. Anolis cblorocyanus, A. coeles- all samples except that from Tortue, there tiniis. A. cybotes, and A. distichus as food are ample numbers of individuals. There items in JJ. catesbyi. He correctly con- are no color data available for specimens in sidered these prey animals as inhabiting collections other than the ASFS, and even trees, tree trunks, and bushes, and stated that in my own material I do not have sufficient U. catesbyi "takes arboreal food almost ex- information to show if differences in dorsal clusively." green hue are individual or populational Uromacer catesbyi occurs from below sea variations. In specimens from other col- level (various localities in the vicinity of lections, it is often difficult to determine Etang Saumatre and Lago Enriquillo in the whether the pale lateral line was present, Cul de Sac-Valle de Neiba plain ) to eleva- since these snakes become readily discolored tions of 5000 feet ( 1525 meters) at Peneau after improper preservation, and many long- on the Montagne Noire in Haiti. preserved snakes are now black and without The abundance of U. catesbyi varies on any pattern indications. However, it seems the satellite islands. This is remarkably dem- reasonable that, if all freshly collected ma- onstrated by contrasting the situation on terial from a region shows certain pattern Ile-a-Vache with that on He de la Tortue. features, these features were also present On the former, during two 1-day visits with in older material, ample native assistance, we secured 59 U. Schlegel's (1837:226-227) description of 134 Tulane Studies in Zoology and Botany Vol. 16 Dendrophis catesbyi is brief but sufficiently detailed to allow the name to be positively associated with one population of the species. His data ( 170 ventrals and 184 subcaudals; uniform grass green above, bluish green below, the two colors separated by a whitish line bordered above by black on the sides of the head ) show that the specimen before him almost certainly was a female from the Tiburon Peninsula population. Since col- lection of the ;>nak.e was attributed to Alex- andre Ricord, a Frenchman who likely col- lected in Haiti ( which at that time was the French colony of St.-Domingue ) and who most probably collected in the vicinity of one of the prominent settlements such as Port-au-Prince, where snakes with similar scale counts and pattern occur, the appli- cation of nominate status to the Tiburon snakes is affirmed. Systematic Accounts Vromacer catesbyi Schlegel D.iendropbis ) catesbyi Schlegel, 1837, Essai sur la physionomie des serpents, 2:226. (Type locality — St.-Domingue). Uromacer catesbyi Dumeril and Bibron, 1854, Erpet. gen., 7:721. Uromacer scandax Dunn, 1920, Proc. New England Zool. Club, 7:43. (Type locality — "Tortuga Island" ). Description: A moderately long (males to 718 mm, females to 908 mm snout-vent length) and slim colubrid snake with 17 or 19 (infrequently 21 ) scale rows behind the head, 17 scale rows at midbody, and 11 scale rows before the vent; 157 to 177 ventral scales in males, 155 to 179 in females; dorsal scales smooth and without apical pits; head scutellation of the normal unspecialized colu- brid type, although the snout is slightly elongate, with modally 11 loreals, 11 pre- oculars, 2/2 postoculars, and 1 + 2/1 + 2 temporals; supralabials usually 8/8, infra- labials usually 10/10; anal divided; sub- caudals paired, 172 to 208 in males, 159 to 201 in females; dorsum green, the individual scales usually not conspicuously outlined with black along their free margins; lower sides with or without a sharply contrasting longitudinal line, white, pale green, or blue, embracing scale rows 1 to 3 or any combi- nation thereof; venter immaculate pale green, this color paler than the dorsum, often ex- tending dorsally onto the lower two or three dorsal scale rows and, in the absence of a pale longitudinal lateral line, blending into, or sharply set off from, the darker dorsal green color; a dark green to black mask-like line from the snout through the lower half of the eye onto the temporal region, bounded below by the pale supralabial coloration, and extending onto the neck where it rapidly disintegrates; top of head immaculate green, occasionally ( by population ) with scattered irregular darker green markings or blotches; hemipenis short ( extending about the length of 5 or 6 subcaudals), deeply bifid, the sulcus spermaticus bifurcating to send a branch along each of the distal forks of the hemipenis; the sulcus spermaticus traverses an area which is finely papillose or spinose after first coursing across a prominent me- dian basal lobe or ring which is densely set with minute spines; the non-sulcate surface is laterally covered with closely set enlarged spines which stop abruptly at the papillose region bordering the sulcus, and which more medially are sharply set off from a more or less cordate area, its apex directly basally, of minute spines; this cordate area is ex- panded distally to form almost a pair of apical discs which form the non-sulcate sur- face of the distal hemipenial forks. (De- scription based on the everted organs of ASFS V9565 and ASFS V10304). Uromacer catesbyi catesbyi Schlegel, 1837 Definition: A subspecies of U. catesbyi characterized by the combination of very low number of ventral scales (161 to lG9 in males, 156 to 172 in females), high number of subcaudal scales ( 188 to 202 in males, 173 to 190 in females), a prominent sky- blue lateral longitudinal line usually on scale row 2, but often including 1 and 3, upper surface of head immaculate green, and large size. Distribution: The Tiburon Peninsula in southwestern Haiti, east to about the level of Momance ( Figure 1 ) ; specimens from Momance eastward through the Cul de Sac- Valle de Neiba plain and the Peninsula de Barahona in the Republica Dominicana are interpreted as intermediate between the nominate subspecies and subspecies to the north (see discussion beyond). Variation: A series of 18 male and 29 No. 4 Uromacer catesbyi 135 female U. c. catesbyi from the Tiburon Pe- ninsula west of Momance has the following counts ( means and extremes ) ; ventral scales in males 161-169 ( 164.9), in females 156- 172 ( 163.2 ) ; subcaudal scales in males 188- 201 (194.6), in females 173-190 (182.1); supralabials usually 8/8 (45 snakes) with 2 snakes having 7 8, infralabials usually 10 10 (40 snakes) with aberrant counts of 8 8 (1), 9/10 (3), 9 11 (1), and 10/11 (1); loreal scales 1 1 ( ASFS X3059 with 0 0 loreals), preocular scales 11 (CM 37788 with 2 2 preoculars), postocular scales 2/2 (MCZ 70138 with 2 3 posto- culars), temporals 1 + 2/1 + 2 (CM 37788, MCZ 70135, ASFS X3213 with 1 + 1 bi- laterally; MCZ 70138, MCZ 64798 with 1 + 1 unilaterally ) ; largest female ( MCZ 65207 ) with a snout-vent length of 830 mm, largest male (MCZ 70131) 685 mm. Material from the Port-au-Prince region, where Ricord may have secured the holotype, is intermediate between the nominate sub- species and material from further north; snakes from a short distance west of Port-au- Prince (Momance) clearly resemble those from more distal portions of the peninsula. Even in the Port-au-Prince region, some snakes have the typical pale blue lateral line of U. c. catesbyi; this line fades to white in long-preserved snakes and I have little doubt that the holotype possessed this pattern fea- ture while alive. In ventral and subcaudal counts, the holotype agrees with females from both the Tiburon Peninsula ( if slight differences in techniques of counting both complements of scales are taken into con- sideration ) and the Port-au-Prince region. It is of course possible that the holotype originated from some more western locality on the peninsula; sites such as Jeremie and Les Cayes have long been prominent sea- ports for this region. There is little doubt, however, that U. c. catesbyi is the proper name for the Tiburon Peninsula snakes. Specimens from Camp Perrin were dark green ( PI. 22 K 4 to PI. 24 C 7; all color designations from Maerz and Paul, 1950) in life, with both supra- and infralabials pale green. There was a pale blue (PI. 25 A 4) longitudinal line of scale row 2, in some specimens including portions of rows 1 and 3, often prominent only anteriorly but in some snakes continuing posteriorly to the vent. The venters ( including the chin ) were pale yellow-green (PI. 17 H 5 to PI. 17 C 4), this paler coloration extending dor- sally onto at least scale row 1 , tliereby giving a tricolor appearance to the lower sides — dark green, pale blue, and yellow-green, from dorsal to ventral. The iris was gold above and below, and brown anteriorly and pos- teriorly, thus giving a quadripartite effect; the black mask was outlined above with faintly paler green. Horn (1969) commented on the occur- rence of a gray morph of V. catesbyi near Miragoane. His data were based upon ex- amination of five snakes in the Harvard col- lection, of which the number of only one (MCZ 66350) was reported. In actuality, there are six U. catesbyi in the MCZ collec- tion (25555, 25556, 66349, 66350-51, and 66352) from the Miragoane region, and another in the Carnegie Museum (CM 37875). 1 had examined all these specimens prior to the appearance of Horn's paper and considered them in no way different in pig- mentation of pattern from other preserved snakes from the Tiburon Peninsula. I have re-examined them in the light of Horn's contention that they were gray above and cream below in life and find that as pre- served they are bluish to greenish blue above and pale grayish green below — all colors which occur in preserved U . catesbyi from throughout its range. Since green is an es- pecially fugitive color in preservative, and since the precise color of the preserved snake depends upon the original fluid in which it was preserved and the length of time it was stored in the fluid prior to being trans- ferred to the appropriate concentration of alcohol for permanent storage, I do not feel that these snakes ( among which must be the four remaining snakes which Horn ex- amined) differed in any strongly chromatic manner from other snakes assigned to U. c. catesbyi. The unproved dichromatism in U. catesbyi needs further confirmation before it is established. Scale counts of all subspecies are grouped in Table 1 for ready reference. The various subspecies comparisons are grouped after the subspecies accounts, and this section is in turn followed by a discussion of the prob- able history of the species on Hispaniola. Specimens examined: Haiti. Di'pt. dii Sud, Jeremie, 2 (MCZ 3603); Fond Rouge Daye, nr. Jeremie ( not mapped ) ( MCZ 136 Ttdane Studies in Zoology and Botany Vol. 16 70128-33 ) ; Place Negre, nr. Jeremie (MCZ 64798-800); Tosia, nr. Jeremie (not mapped) (MCZ 70138); Paroty, nr. Jeremie ( not mapped ) ( MCZ 70137 ) ; Mayette, nr. Jeremie (MCZ 70135-36); Lantinzi, nr. Jeremie (not mapped) (MCZ 70134); Ri- verdi, nr. Jeremie ( not mapped ) ( MCZ 70127); frou Bois (MCZ 74521-22); Camp Perrin ( ASFS X2976, ASFS X3059- 65, ASFS X3153-57, ASFS X3252-53); Petit Trou de Nippes ( USNM 80812); nr. Etang Miragoane (MCZ 25556); Dept. de I' 0 nest. Bascap-Rouge, 10 km NE Jacmel ( CM 37788-89, MCZ 65207-08 ) ; Momance (MCZ 8670); either Dept. de I'Ouest. or Dept. dii Slid. Commune Aquin, nr. Mi- ragoane (CM 37875, MCZ 66349); 6 mi. ( 12.6 km) from Miragoane (MCZ 25555); Butete, nr. Miragoane (MCZ 66350); Min- grette, nr. Miragoane (MCZ 66352). Uromacer cateshyi iiisulaevaccariini,^ new subspecies Holotype: CM 45875, an adult female, western end, Ile-a-Vache, Dept. du Sud, Haiti, one of a series collected by native col- lector on 4 August 1962. Original number ASFS X3539. Paratypes: ASFS X3381-87, ASFS X3537-38, ASFS X3540-46, same data as holotype; AMNH 103215-24, CM 45877- 86, MCZ 92076-85, USNM 165937-48, same locality as holotype, native collectors, 6 August 1962. Definition: A subspecies of U. catesbyi characterized by the combination of low number of ventral scales ( 162 to 173 in males, 159 to 172 in females), moderate to high number of subcaudal scales ( 180 to 204 in males, 172 to 200 in females), a pale greenish lateral longitudinal line on scale row 2, occasionally involving a portion of scale row 3, upper surface of head immacu- late green, and moderate size. Distribution: Ile-a-Vache, off the south- western coast of the Tiburon Peninsula, Haiti ( Figure 1 ) . Description of holotype: An adult female with the following scale counts: ventrals 169, subcaudals 196, supralabials 8/8, infra- labials 10/10, loreals 1, 1, preoculars 12, ^ From insula (island) and racca (cow), an allusion to the distribution of the subspecies, Ilc- a-Vache. postoculars 2/2, temporals 1 + 2/1 + 2; dor- sal scale row formula 19-17-11; snout-vent length 712 mm, tail 595 mm. In life, dorsum green (PL 22 L 4), with a pale greenish anci faint longitudinal line on scale row 2; venter pale yellow-green with a faint yellow line below the dark green mask-like line; iris golden above, black anteriorly and posteriorly, and dark brown below. Variation: The series of 25 male and 34 female f7. c. insulaevaccarufn has the follow- ing counts ( extremes and means ) : ventral scales in males 162-173 (167.0), in females 159-172 ( 166.3); subcaudal scales in males 180-204 (192.3), in females 172-200 (184.9); supralabials usually 8/8 (56 snakes) with one snake having 7/8 and two snakes having 9/9, infralabials usually 10/ 10 (31 snakes) with aberrant counts of 9/10 (2), 10 11 (15), 11/11 (9), and 11/12 (1); loreal scales 1 '1 (AMNH 103215 with 0/0; AMNH 103222 with 0/1), preocular scales 1 1 (holotype and USNM 165946 with 12), postocular scales 2/2 (AMNH 103220 with 2/3), temporals 1 + 2/1 + 2 (ASFS X3386 with 2 + 2 bi- laterally; MCZ 92078, USNM 165946, with 1 + 1 unilaterally ) ; largest female ( AMNH 103219) with snout-vent length of 800 mm, largest male (ASFS X36l6) 615 mm. All specimens collected by myself and parties in Ile-a-Vache agree with the defi- nition of the subspecies as far as color and pattern are concerned. One individual had the lateral longitudinal line pale blue like nominate catesbyi rather than pale green. The dorsal coloration in the series varied from the dark green of the holotype ( PI. 22 L 4 ) to a paler hue ( PI. 2 1 J 5 ) . Remarks: U. c. insulaevaccarum occupies Ile-a-Vache, an island separated from the southern coast of the Tiburon Peninsula of Haiti by a channel 10 kilometers in width. The Ile-a-Vache fauna has several endemic subspecies of reptiles ( Diploglossus costatus nesobous, Anolis distichus juliae, Anolis coelestinus pecuarius, Ameiva taeniura aecju- orea, Dromicus parvifrons rosamondae) , but other reptiles and one amphibian ( Eleuther- odactylus pictissimus, Diploglossus stenurus, Leiocephaliis melanochloriis ) whose variation has been studied in detail, do not show any local differentiation. Perhaps the latter group No. 4 Uromacer catesbyi 137 4-* .3 !1) ID c « a 4--^ C/3 > a c/^ a; CL, ^ o 2 ;^ :«5 -5 -5 l-O O lO o o o 00 CO ^ o ^ ^ CD CO CD 00 CD -f ICO CD t^ OO xic CD t^ CD t^ o lo o in o -t t-io CO CO lOlO t- CD CO CO c^ lO CO t^ CD CO CO coi^ LO ^ COCO CO X CO ^ LO CO CO .6(349- .5(336- 5(341- 4(339- 1 CO o 3(356- 0(348- 1(338- 2(332- X LO CO CD ' 1 1 CO ^ CO CO 02 rf lO ~V CO CO 02 r-H lOlO coco CO -V CO CD LO CO CO T^ CO coco CD LO t V CO CO o CD CO X— 1 CO CO CO CO i-H O -t< O O 02 o o CM ^ Ol X CD CD •—i .—1 J^ci CD in ^H T— 1 Ol 02 CD LO ^H 1— ( Ol d, CD CD Ol t- CD CD -H CO CD CD 1—1 ^H t^ lO CD CD ■— I r— 1 oq CD CD T— 1 ^H X Ol O CO Ot- CD-IH ^ X -t o LO cot- '^ CO CD CD t- CD CD CD CD LO CD CD l>t> t- CD CD CD ox t^ CD Ol LO CO CO CD CD X02 lO^ r-H Ol Ol CO CD CO CD O cob- ■— I X -o Ot I— I Ol CO t- ►a Slj 138 Tulane Studies in Zoology and Botany Vol. 16 represents more recent arrivals than does the former. In addition to the specimens herein desig- nated paratypes, there is a long series of JJ. catesbyi in the collection of the Museum of Comparative Zoology. Most of our snakes were collected by natives; one was secured in my presence by a boy who took the snake from the crown of a coconut palm with a sloping trunk, which the snake had just as- cended with great rapidity. Uromacer catesbyi cereolineatus," new subspecies Holotype: MCZ 92074, an adult male, vicinity of Pointe Sable, He Grande Caye- mite, Dept. du Sud, Haiti, one of a series collected by native collectors on 18 March 1966. Original number ASFS V9563. Paratypes: ASFS V9561-62, ASFS V9564- 66, CM 45887-88, same data as holotype. Associated speciiuen: Haiti. Dept. du Sud, He Petite Cayemite, 1 ( USNM 80825). Definition: A subspecies of U. catesbyi characterized by the combination of low number of ventral scales (162 to 170 in males, 160 to 169 in females), high number of subcaudal scales ( all males with incom- plete tails; 177 to 185 in females), a white lateral longitudinal line on scale row 2, and upper surface of head immaculate green. Distribution: He Grande Cayemite (and presumably He Petite Cayemite), off the northern shore of the Tiburon Peninsula, Dept. du Sud, Haiti ( Fig. 1 ) . Description of holotype: An adult male with the following scale counts: ventrals l68, subcaudals 173+ (tail incomplete), supralabials 8 8, infralabials 10/10, loreals 0/0, preoculars 11, postoculars 2/2, tem- porals 1 + 2/1 + 2; dorsal scale row formula 17-17-11; snout-vent length 640 mm, tail 558 mm, incomplete. In life, dorsum dark green (PI. 24 H 4), with a prominent white lateral longitudinal line on scale row 2; venter yellow-green; chin and throat very pale green, nearly white; mask-like line bordered above with pale yellow-green. Iris color not noted. Variation: The series of six male (in- cluding one male from Petite Cayemite) and three female U. c. cereolineatus has the fol- lowing counts (extremes and means): ven- tral scales in males 162-170 (166.0), in females 160-169 (165.7); total subcaudal scales unknown in males (in three males with nearly complete tails subcaudal counts vary between 173 and 194), in females 177- 185 (181.0); supralabials always 8 8, in- fralabials usually 10 10 (6 snakes) with aberrant counts of 10 11 in two snakes; loreals IT (holotype with 0 0), preocular scales 11, postocular scales 2 2 (ASFS V9565 with 12), temporals 1 + 2 1+2; only female which can be reliably measured (CM 45888) has a snout-vent length of 410 mm, largest male ( holotype ) 640 mm. Color in life (Grande Cayemite specimens only; data taken by Richard Thomas ) varied from dark to light green ( PI. 24 H 4, PI. 23 H 2, PI. 22 I 4 ), and the lateral longitudinal lines were white in both sexes. In two spec- imens the white lateral lines faded out on the posterior third of the body, but in other in- dividuals the lines persist to the vent. The venters in the series were pale green to yellow-green, and the chin and throat were very pale green (almost white). Uromacer catesbyi hariolatus.'' new subspecies Holotype: USNM 165936, an adult male, 2 mi. (3.2 km) W Trou du Nord, Dept. du Nord, Haiti, taken by Richard Thomas on 8 April 1966. Original number ASFS V10214. Paratypes ( all from Dept. du Nord, Haiti):' ASFS V10168, 1 mi. ( 1.6 km) E Terrier Rouge, native collector, 8 April 1966; ASFS VI 0303, Port Margot, native collector, 10 April 1966; ASFS V10304, 5 mi. (8.0 km) E Limbe, E. Cyphale, 10 April 1966; ASFS V10305, Cap-Haitien, native collector, 10 April 1966; MCZ 37604, MCZ 37606, Cap-Haitien, 31 March 1934, Utoivana ex- pedition. Associated specimens: Haiti, Dept. du Nord Quest, Mole St. Nicholas (MCZ 64797, MCZ 62697 ) ; Bombardopolis ( MCZ 62694-96); Dept. de I'Artibonite. St. Marc (AMNH 49767); Hinche (MCZ 25553- 54); Dept. de I'Ouest, Mirebalais (MCZ "From cereus (waxen) and linea (a thread), '"'From harolior (to predict), an allusion to the an allusion to the white lateral longitudinal line. expected occurrence of a north island subspecies. No. 4 Uromacer catesbyi 39 68534); Duvier, nr. Mirebalais (not mapped) (MCZ 68538); Lancironelle, nr. Mirebalais (not mapped) (MCZ 68536-37); Ledie, nr. Mirebalais (not mapped) (MCZ 68543); Fer-a-cheval, nr. Mirebalais (MCZ 68535); Boucan, nr. Mirebalais (not mapped) (MCZ 68539-42); 2.1 mi. (1.9 km) NE Barrage de Peligre (ASFS X2214). Definition: A subspecies of U. catesbyi characterized by a combination of high to very high number of ventral scales ( 162 to 177 in males, 167 to 176 in females), mod- erate to high number of subcaudal scales (183 to 203 in males, 177 to 194 in females), no pale lateral longitudinal line but lower sides grading gradually ( on the first three scale rows ) from yellowish green venter to dark green dorsum, upper surface of head immaculate green, and moderate size. Distribution: Haiti, north of the Cul de Sac Plain; intergradient specimens {cates- byi X hariolatus ) from the vicinity of Port- au-Prince and the extreme western Cul de Sac ( Figure 1 ) . Description of bolotype: An adult male with the following scale counts: ventrals 177, subcaudals 183, supralabials 8 8, infra- labials 10, 10, loreals 1 1, preoculars 11, postoculars 2/2, temporals 1 + 2/1 -f 2; dor- sal scale row formula 17-17-11; snout-vent length 610 mm, tail 475 mm. In life, dorsum dark green, venter yel- lowish green, the ventral color grading grad- ually on the lowermost three scale rows into the dorsal color, without the interposition of a pale longitudinal lateral line; chin, throat, and supralabials pale yellowish green; mask-like line bordered above by pale yel- lowish green. Iris color not noted. Variation: The series of 16 male and 10 female U. c. hariolatus has the following counts (extremes and means) : ventral scales in males 162-177 ( 171.6), in females 167- 176 ( 171.4); subcaudal scales in males 183- 203 (191.6), in females 177-194 (183.8); supralabials usually 8 8 (23 snakes) with variation of 7 7 ( 1 ), and 7/8 ( 1 ), and 8 9 ( 1 ); infralabials usually 10 10 (24 snakes) with aberrant counts of 9 10 ( 1 ) and 10 11 (1); loreal and preocular scales 11 in all specimens, postocular scales 2 2 (ASFS X2214 with 2 4, ASFS VI 0305 with 2 3), temporals 1 + 2/1 + 2 (MCZ 62697 and ASFS V10168 with 1 + 1 unilaterally; MCZ 68538 with 1 + 1 bilaterally); largest female (MCZ 68539) with snout-vent length of 790 mm, largest male (MCZ 68538) 645 mm. All specimens were presumably some shade of green in life. The snake from Ter- rier Rouge was recorded as having the dor- sum PI. 29 A 10 and the venter PI. 17 C 8 in life, whereas the Port Margot snake had the dorsum about PI. 2 1 I 10. The lateral scale rows ( 1-3) were paler than the balance of the dorsum, and showed a gradation be- tween the ventral color and that of the back. One snake ( MCZ 64797 ) from Mole St. Nicholas on Haiti's Presqu'ile du Nord- Ouesr, is imique among the hariolatus series in that the lower sides are sharply demar- cated from the dorsum ( but there is of course no longitudinal lateral line). This style of sharply contrasting lower sides agrees better with Dominican IJ . catesbyi (see dis- cussion following ) rather than with other hariolatus. Three other specimens from the Presqu'ile du Nord-Ouest (Mole St. Nicho- las and Bombardopolis ) are typical hari- olatus in this feature, however. Uromacer catesbyi pampineiis,^ new subspecies Holotype: MCZ 92075, an adult female, 2.1 mi (3.4 km) N Hato Mayor, El Seibo Province, Repiiblica Dominicana, taken by Ronald F. Klinikowski on 18 June 1963. Original number ASFS X7875. Paratypes ( all from the Repiiblica Do- minicana ) : ASFS V637, 3 km NE Centro de Boya, San Cristobal Province, R. Thomas, 22 August 1963; ASFS VI 118-19, 2.5 km NW Boca de Yuma, La Altagracia Province, native collector, 4 September 1963; MCZ 16323-24, La Romana, La Romana Province, E. Lieder, 1922; ASFS V2469-70, 5.1 km E. Santo Domingo, Distrito Nacional, D. C. Leber, R. Thomas, 19 June 1964; MCZ 79297-60, Central Ozama, Distrito Nacional, J. D. Lazell, Jr., 29 December 1963; MCZ 57987, Santo Domingo zoo, Distrito Na- cional, J. A. Rivero, 29 March 1953. Associated specimens: Repiiblica Domin- icana. Monte Cristi Province. 2 km NE Palo Verde, 5 (ASFS V1320-24); 1 km S Palo Verde, 1 (ASFS V1351); Laguna de Salo- dillo, 7 km SE Pepillo Salcedo, 2 ( ASFS * From pampiiieiis (pertaining to vine tendrils or leaves), an allusion to the green coloration. 140 Tulane Studies in Zoology and Botany Vol. 16 o (M CO 00 ^. ^-2 1 o o = c o a « 3 (— ' X ^ ^ n -= ^ Q ■■^fe ^ Qj r- -*-' O r^ •4-> ^ o =^ ^.f, *; ;»; ^ c^ = ?:^ =- ^ 5j Oj 5j ^> rt ^ ;^ OJ > r^ 1— I C/} s %.. ij « ^ o Vj U ^ ^ o u '"'i - > -|a ■2 !^ CD a 5j 05 Co r- aj a ■ o H'^ S ,,_G Oj ■ 2 S' ^ ^ co-.ii S „ 2 O > c s a a C ~ ^ /^ '^ ^ .G a U 0) a; o 0; ».G N) ** g C (U 0 °^ i a bC 0,0-" 1>> ;y: a a) rt bc G C ' I I ^riG O .2 a cs u ti) 'a 2i ^ 5 c .^ "" a; G ^ O ° G '^ « C/2 No. 4 Uromacer catesbyi 141 V1458, ASFS V1423); Dajab6n Proimcc, 1 km S Loma de Cabrera, 900 feet (275 me- ters), 1 (ASFS VI 173); Puerto Plata Prov- ince, Sosua, 1 (MCZ 43662); Choco, 1 (MCZ 13676); 8 km N Pena, 1 (MCZ 57763); Samand Province. Sanchez, 1 (CM 8099); La Vega Province, 1 mi. (1.6 km) WSW Constanza, 4000 feet (1220 meters), 1 ( ASFS X8731 ) ; 12 km NE Jarabacoa, 2000 feet (610 meters), 2 (ASFS VI 949, ASFS V14152); 14.4 km E La Vega, 1 (ASFS V4214); La Estrelleta Province, Cerro de San Francisco, 3 km E Banica, 1 (MCZ 57767). Definition: A subspecies of U. catesbyi characterized by the combination of mod- erate to high number of ventral scales ( l6l to 174 in males, l63 to 171 in females), low number of subcaudal scales ( 172 to 189 in males, 162 to 183 in females), no pale lat- eral longitudinal line but lower sides blue- green to yellow-green on lowermost three scale rows, changing abruptly to green of dorsum, upper surfaces of head immaculate green, and small size. Distribution: The Republica Dominicana north of the Valle de Neiba ( Figure 1 ) . Description of holotype: An adult female with the following scale counts: ventrals 164, subcaudals 168 (tail slightly incom- plete), supralabials 8 8, infralabials 10/10, loreals 1/1, preoculars 1/1, postoculars 2/2, temporals 1 + 11 + 2; dorsal scale row for- mula 17-17-11; snout-vent length 722 mm, tail 529 mm. In life, dorsum green (PI. 29 F 11 ); first two scale rows, plus half of third scale row, pale blue-green (PI. 25 B 7); venter and supralabials pale green (PI. 17 H 8); mask- like line dark green, bordered above by faintly paler green. Iris golden brown above, brown below. Variation: The series of 13 male and 17 female U. c. pampineus has the following counts (extremes and means) : ventral scales in males 161-174 (167.4), in females l63- 171 ( 166.8); subcaudal scales in males 172- 189 (179.8), in females 162-183 (171.1); supralabials usually 8 8 ( 29 snakes ) with one snake having 7 8, infralabials usually 10/10 (25 snakes) with aberrant counts of 9^9 (1 ), and 10 11 (3); loreal scales 1/1 ( MCZ 79260 with 0 1), preocular scales 1/1 (ASFS VI 118 with 2 1, ASFS V4214 with 2/2), postocular scales 2 2 (ASFS VII 73 with 12), temporals 1+2 1+2 (holotype and ASFS V1458 with 1 + 1 uni- laterally, MCZ 79257 with 1 + 1 bilaterally, ASFS V637 with 1 + 3 unilaterally ) ; largest female ( CM 8099 ) with snout-vent length of 770 mm, largest male (ASFS V1321) 610 mm. All specimens were green in life; a female from near Jarabacoa was recorded as being PI. 19 D 9 above. The venters were uniformly a paler yellow-green (recorded as PI. 25 A 7 in the Jarabacoa snake ) ; the paler coloration extended onto the lowermost two or three dorsal scale rows as a distinct longitudinal pale area. In some individuals, such as the holotype, the lower sides were pale blue- green, in contrast to both the yellowish venter and the more pure green dorsum. In preserved specimens the lower sides are sharply bicolor in may well-preserved snakes; in other individuals longer preserved or discolored, the bicolor condition of the sides is no longer discernible. Remarks: No certain intergrades between hariolatus and panipineus are known from along Dominico-Haitian border. Specimens from localities near this border (Monte Cristi, Dajabon, and La Estrelleta provinces) include snakes which have low subcaudal counts. A male from Monte Cristi (ASFS V1320) has a subcaudal count of 189, the upper extreme of male painpineus: the total underbody scales ( ventrals subcaudals ) of this snake are 357, at the point of overlap of this count between pampineus and hari- olatus. Perhaps the Pepillo Salcedo region is one of extreme intergradation between pampineus and hariolatus, with the popula- tion more closely resembling the Dominican snakes than the Haitian. The snake with the highest female subcaudal count is from La Estrelleta Province; this snake has 183 subcaudals and a total underbody count of 347, the upper extreme of pampineus fe- males. It too probably shows the genetic influence of hariolatus upon a predominantly pampineus population, although with but a single specimen details are lacking. U. c. pampineus is widespread throughout the Republica Dominicana and occurs from sea level to elevations of at least 4000 feet (1220 meters) in the Vallc de Constanza in the Cordillera Central. The holotype was collected in a Theobroma grove, and one snake from near Jarabacoa was collected as 142 Tulane StJidies in Zoology and Botany Vol. 16 it climbed about the lower branches of a deciduous tree in what is essentially upland (610 meters) pine woods. The specimen from Centro de Boya was taken as it crawled across a termitarium 4.5 meters above the ground, and the long series from Palo Verde was collected on a barbedwire fence adjacent to anole-rich lowland woods. Uromacer catesbyi frondicolor,'' new subspecies Holoiype: MCZ 93162, an adult male, Degoute, He de la Gonave, Haiti, one of a series collected by George Whiteman in December 1965. Paratypes (all from He de la Gonave): MCZ 93163-65, same data as holotype: ASFS X2431-32, Etroits, E. Cyphale, 17 July 1962; ASFS X3322, Etroits, E. Cyphale, 26 July 1962; USNM 75926, Anse a Galets, J. S. C. Boswell, 21 September 1928; MCZ 93171, Platon Yeye (not mapped), G. Whiteman, December 1965; MCZ 93138, Boisederet (not mapped), G. Whiteman, December 1965; MCZ 93167-70, Ravine Fogue (not mapped), G. Whiteman, De- cember 1965; MCZ 93172-73, Bois Boule (not mapped), G. Whiteman, December 1965; MCZ 80830-31, Ti Palmiste, 6 km from Pointe-a-Raquettes, G. Whiteman, sum- mer 1964; USNM 63115, "Gonave Island," W. L Abbott, 16 March 1920; MCZ 12869, "La Gonave," G. M. Allen, 1918. Definition: A subspecies of U. catesbyi characterized by a combination of moderate to high number of ventral scales (167 to 175 in males, 165 to 171 in females), (mod- erate ? to) very high number of subcaudal scales ( 192 to 208 in males, 175 and 180 in two females), no pale lateral longitudinal lint, the lower sides grading gradually (on the first three scale rows) from the light green venter to the dark green dorsum, upper surfaces of head immaculate green, and mod- erate size. Distribution: He de la Gonave, Haiti. Description of holotype: An adult male with the following scale counts: ventrals 171, subcaudals 193, supralabials 8 8, infra- labials 10/10, loreals 1^1, preoculars 1/1, postoculars 2/2, temporals 1 + 11 + 3; dor- ■' From frons (leaf) and color (color), an al- lusion to the green color. sal scale row formula 18-17-11; snout-vent length 688, tail length indeterminate because of damaged tail. Preserved, the holotype is blue-green above, slightly paler below. The supralabials are concolor with the venter, and the black mask-like line is prominent and bordered above on the temporal region by pale bluish green. Iris color indeterminable. Variation: The series of 1 1 male and 8 female U. c. frondicolor has the following counts (extremes and means): ventral scales in males 167-175 ( 170.4), in females 165- 171 ( 168.0); subcaudal scales in males 192- 208 ( 197.0), in two females with complete tails 175-180 (177.5); supralabials 8/8 in all specimens, infralabials usually 10/10 (11 snakes) with aberrant counts of 9 'TO (3), 10/11 (3), 11/11 (2) and 1112 (1); loreal scales IT, preocular scales usually 11 (MCZ 93168 and MCZ 93165 with 12), postoculars 2 2 (MCZ 93138 with 1 /2 ), temporals 1 + 2/T + 2 (ASFS X3432, MCZ 12869, MCZ 93138 with 1 + 1 uni- laterally; MCZ 93170 with 1 + 1 bilaterally; holotype with 1 + 1 T + 3 ); largest female ( MCZ 80830 ) with snout-vent length of 755 mm, largest male (holotype) 688 mm. The color of the three U. c. frondicolor seen in life was recorded as light green below and dark green above, and there was no blue lateral line. The remaining paratypes have been preserved for various periods, and little can be said about the coloration and pattern in life of this subspecies. Remarks: The relatively few specimens of U. c. frondicolor available suggest that the species is uncommon on He de la Gonave. One of the specimens from Etroits was taken by natives in a lime grove. Since so few of the localities for U. c. frondicolor are pre- cisely locatable, the altitudinal distribution of the subspecies is unknown; however, Etroits is on the coast in an extremely xeric situation and Ti Palmiste lies inland at an elevation of about 400 meters. Uromacer catesbyi scandax Dunn, 1920 Definition: A subspecies of U. catesbyi characterized by a combination of high to very high number of ventral scales (172 in single male, 172 to 179 in two females), high (?) number of subcaudal scales (187 in female with complete tail), no pale lateral No. 4 Uromacer catesbyi 143 longitudinal line and, apparently, lower sides not set off chromatically from the upper dorsum, and upper surface of head with scat- tered darker areas (in life green? ) on a paler ground at least in juveniles. Distribution: He de la Tortue, off the northern Haitian coast ( Figure 1 ) . Variation: A single male JJ. c. scandax has 172 ventrals, subcaudals indeterminate, 8/8 supralabials, 10 10 infralabials, 1 1 loreals, 1 1 preoculars, 2 2 postoculars, 1 + 2/1 +2 temporals, dorsal scale row formula 19-17-11, snout-vent length 540 mm; two females have 172 and 179 ventrals (mean 175.5 ), 187 subcaudals in one specimen, 8/8 supralabials, 10/10 infralabials, 1/1 loreals, 11 preoculars, 2 2 postoculars, 1 -f 2/1 + 2 temporals, 19-17-11 and 17-17-11 dorsal scale rows; one female has a snout-vent length of 525 mm and a tail length of 410 mm. The status of the Tortue population of U. catesbyi remains equivocal. Dunn (1920) described U. scandax from a single female from "Tortuga Island," and Cochran (1941: 334) accepted this form as a species distinct from U. catesbyi. Cochran also (op. cit.: 332-333) considered two other specimens from Tortue as V. catesbyi, thereby implying that two short-snouted species of JJroniacer occurred on that island. Mertens ( 1939:78), however, considered scandax only subspe- cifically related to U. catesbyi. I have studied the three specimens available to Cochran and can see no reason either to consider scandax a species distinct from catesbyi or to exclude the two other Tortue specimens from scandax. These three specimens, con- sidered together, seem to indicate that scan- dax is a moderately well defined subspecies of U. catesbyi. Lack of an adequate series makes meristic comparisons of scandax with other U. catesbyi subspecies difficult. How- ever, the ventral counts of two female scandax (111 and 179) are at the upper limit of ventral counts for the species (155 to 179), and the count for the single male (172) is high in the total range of this count in male U. catesbyi ( 157-177 ). Except for the high ventral counts in female scandax. the next highest counts occur in female U. c. hario- latus, which occurs on the adjacent mainland of Haiti. The two smaller specimens of V . c. scandax ( USNM 59920, MCZ 37607 ) are peculiar in that they are now a dull grayish green dorsally and are heavily stippictl with dark gray-green ventrally — a chromatic develop- ment not matched by any other specimens of U. catesbyi. The adult holotype ( USNM 59438) presently is colored like comparably aged specimens. It is likely that fresh speci- mens of scandax will demonstrate chromatic differences, at least in immature specimens, between the Tortue subspecies and those elsewhere. Speci?nens examined: Haiti, lie de la Tortue (USNM 59438— holotype, USNM 59920, MCZ 37607). Uromacer catesbyi inchausteguii,'' new subspecies Holotype: CM 45876, an adult female, environs of Mano Juan, Isla Saona, Repub- lica Dominicana, one of a series collected by native collectors on 28 December 1968. Original number ASFS Vl6l57. Paratypes (all from Isla Saona): ASFS V161 58-59, ASFS V16177, same data as holotype; ASFS V3068, same locality as holo- type, R. Thomas, 19 July 1964; MCZ 92086- 90, same locality as holotype, native collec- tors, 13 August 1968; AMNH 103225-29, USNM 165950-54, same locality as holo- type, native collectors, 14 August 1968; USNM 165949, same locality as holotype, native collector, 29 December 1968; LDO 7-5535, same locality as holotype, native col- lector, 30 December 1968; ASFS V 16321, 2.5 mi. (4.0 km) NW Mano Juan, J. A. Rodgers, Jr., 29 December 1968. Definition: A subspecies of U. catesbyi characterized by the combination of low to very low number of ventral scales (I6O to 167 in males, 160 to 168 in females), very low number of subcaudal scales ( 172 to 176 in males, 159 to 186 in females), no pale lateral longitudinal line, but lower sides (scale rows 1-3) a distinctly paler green than the dorsum, all dorsal scales with a dis- tinct black edging to giwc a definitely squa- mate appearance, upper side of head with dark green markings on a paler green ground, and small size. Distribution: Isla Saona, off the south- '' In honor of Sixto J. Inchaustcuui who was re- sponsible for the opportunity to collect most of the specimens of this subspecies. 144 Tulane Studies in Zoology and Botany Vol. 16 eastern tip of the Republica Dominicana ( Figure 1 ) . Description of bolotype: An adult female with the following scale counts: ventrals 161, subcaudals 163, supralabials 8 8, infra- labials 10/10, loreals 1 1, preoculars 1/1, postoculars 2/3, temporals 1 + 2 1 + 2, dor- sal scale row formula 19-17-11; snout-vent length 680 mm, tail 484 mm. In life, dorsum green, the lowermost three scale rows distinctly paler green than dorsum, a lateral extension of the ventral pale green color; throat and supralabials whitish (very pale green), black mask-like line margined above by very pale green; internasals, pre- frontals, supraoculars, and parietals with dark grayish green flecks, dashes, or rectangular figures (on prefrontals), all very conspic- uous in life. Iris color unrecorded. Variation: The series of 6 males and 17 females has the following counts (extremes and means): ventral scales in males 160- 167 (163.3), in females 160-168 (163.7); subcaudal scales in males 172-176 (174.3), in females 159-186 (167.6); supralabials 8 8, infralabials 10 TO, with two snakes having 9 10; loreal scales 1 1(ASFSV14895 with 0 0), preocular scales I'l, postocular scales 2 2 ( holotype and ASPS VI 4894 with 2 3 ), temporals 1 + 2 T + 2; largest female (ASFS V3068 ) with snout-vent length of 795 mm, largest male ( AMNH 103225) 590 mm. The enitre series of U. c. inchaustegini agrees with the description of the holotype in having the dorsal scales edged with black, these giving a distinctly squamate appearance to the back, and in having flecks, dashes, or small irregular dark figures on the dorsal head scales. The lower sides (scale rows 1-3) are in life often a paler shade of green, con- trasting distinctly with the darker green back, but this feature sometimes is lost after preservation. Reviarks: U. c. inchausteguii appears to be fairly common in the environs of the village of Mano Juan. The specimen taken by Rodgers was secured in open broadleaf forest as it crawled on leaf litter. Of the two species of Uromacer on Isla Saona, U. catesbyi appears to be outnumbered by V. oxyrhynchus, since more specimens of the latter species were brought to us by native collectors. Comparisons Uromacer catesbyi has been shown to be divided into eight subspecies. Three of these subspecies are obviously interrelated more closely than they are to the balance of the subspecies. The subspecies catesbyi, ins/dae- vaccarinn. and cereolineatiis possess a pale lateral line on scale rows 1 to 3, whereas the remaining five subspecies lack this pattern feature. The three lineate taxa differ from each other in the color of the lateral line (pale blue in catesbyi. pale greenish in in- sidaevaccarum, and white in cereolineatiis) . In addition to the differences in lateral line pigmentation, both sexes of V. c. catesbyi have lower mean numbers of ventral scales than do both sexes of inculaevaccarum (male catesbyi 164.8, male instdaevaccarum 167.0; female catesbyi 163.2, female insidaevac- caruni 166.3), although the differences are not striking. In subcaudal scale counts, the situation is anomalous, since male insidae- vaccarum have a lower mean ( 192.3 ) than do male catesbyi (194.6), and female in- sulaeracca/rum have a much higher mean (192.3) than do female catesbyi (182.1). These two subspecies reach approximately the same size. Comparison of catesbyi and insulaevac- carmn with cereolineatiis is made difficult by the short series of the latter subspecies, and the fact that no male cereolineatiis have complete tails. There are also no obviously adult female cereolineatus, so that size com- parisons are not possible. At least male cereolineatiis resemble male insidaevaccarum in maximum known size, rather than there being a resemblance between cereolineatus and adjacent catesbyi in this feature. Total underbody scale counts ( ventrals + sub- caudals ) of female cereolineatus are close, both in means and extremes, to those of catesbyi and are lower than those of insulae- vacarrum. The remaining five subspecies {hariolatus, pampineus, frondicolor, scandax, inchau- steguii) differ as a group from the western subspecies catesbyi-insiilaevaccaru7n-cereolin- eatus series in lacking a lateral line. U. c. hariolatus is characterized by having a grad- ual change from the ventral coloration to that of the dorsum and in having a higher mean number of ventral scales in both sexes in comparison with all the western subspecies No. 4 Uromacer catesbyi 145 (mean in male hariolatiis 171.6, means in males of western subspecies 164.8-167.0; female hariolatus 171.4, means in females of western subspecies 163.2-166.3). U- c. pampineiis resembles hariolatus in being all green and without a lateral line, but hario- latus lacks the strongly contrasting lower scale rows which pa7)ipi)ieiis possesses. The subcaudal counts of hariolatus are consist- ently higher (183-203 in males, 177-194 in females) than in pampineus (172-189 in males, 162-183 in females); the ventral counts of the two subspecies are comparable, with parnpinei/s averaging a few less ventrals than hariolatus. If total underbody scale counts are used to distinguish the two popu- lations, almost complete separation of in- dividuals of both sexes is achieved ( 356 to 375 in male hariolatus, 338 to 358 in male pa?)2pineus: 348 to 361 in female hariolatus, 332 to 347 in female pampineus) . U. c. frondicolor differs from pampineiis in lacking contrastingly colored lower sides and in having a higher number of subcaudal scales (at least in males; only two female frondicolor have complete tails ) . Male pam- pineus, with subcaudal counts of 172 to 189, are completely separable from male frondi- color with subcaudal counts of 192 to 208. Of all the subspecies of U. catesbyi, frondi- color has the highest counts of subcaudal scales. U. c. frondicolor most closely re- sembles adjacent U. c. hariolatus in dorsal and lateral color and pattern. U. c. frondi- color averages less ventral scales in both sexes than does hariolatus ( male frondicolor 170.4, male hariolatus 171.6; female frondi- color 168.0, female hariolatus 171.4), al- though the extremes in male frondicolor are embraced by the extremes in male hariolatus. The situation in subcaudals is anomalous, since the male frondicolor have a higher mean (197.0) than do male hariolatus (191.6); the subcaudal counts of the two female frondicolor on the other hand lie below and at the lower extreme of subcaudal counts in female hariolatus. It seems likely that additional female frondicolor will show that, in this sex also, the subcaudal count has both a higher mean and upper extreme than in female hariolatus. The remaining two subspecies, scandax and inchausteguii. differ from all others in that both have dark cephalic markings; these markings occur apparently only in juveniles in scandax and in all specimens in inchau- steguii. Although scandax is represented by only three specimens, the ventral counts from these three snakes are at the upper ex- treme ( females ) or high ( male ) for the species. On the other hand, inchausteguii has the lowest mean number of subcaudal scales and a low mean number of ventral scales. Total underbody scale counts in in- chausteguii are 332 to 343 in males, 319 to 350 in females. Male inchausteguii are com- pletely separable from male catesbyi, frondi- color and hariolatus. when this count is used as an index, and there is very little overlap between inchausteguii and male insulaevac- carum and pampineus. As far as females are concerned, there is little overlap between total underbody counts of inchausteguii and hariolatus (348-361). The darkly margined scales of inchausteguii differentiate that sub- species from all others. Discussion Uromacer catesbyi. as defined here, is comprised of eight subspecies, three of which occur on the main island ( catesbyi, hariolatus, pampineus ) and five on satellite islands (insulaevac carum, cereolineatus, fron- dicolor, scandax, inchausteguii) . The distri- bution of the subspecies in U. catesb)i is similar to that of Drotnicus parvifrons Cope (see Thomas and Schwartz, 1965); in that colubrid snake, nine subspecies are recog- nized, four of which occur only on the main island, four on the satellites, and one on both a satellite island and the adjacent mainland. Details of subspecific boundaries in U. cates- byi and D. parvifrons on Hispaniola are not comparable, although a distinct subspecies of each occurs on the Tiburon Peninsula. I have examined only a few specimens of U. catesbyi from the Peninsula de Samana in the northeastern Repiiblica Dominicana. This peninsula, with its narrow swampy neck, has been a center of differentiation for several species of amphibians and reptiles ( Eleutherodactylus weinlandi, Diploglossus stenurus. Droniicus parvifrons); it is possible that still another subspecies of U. catesbyi inhabits the Peninsula de Samana. No mention has thus far been made of the complexities of the interrelationships be- tween the subspecies catesbyi, hariolatus, and pampineus in, and south of, the Cul de Sac- 146 Tulane Studies in Zoology and Botany Vol. 16 Valle de Neiba plain which separates the old north and south (sens// Williams, 1961 ) islands of Hispaniola, and on the Peninsula de Barahona. A discussion of the problems in this region, and the interpretation of these problems is now presented. The region of intergradation between nominate catesbyi and hariolatns is in the vicinity of Port-au-Prince and the western extreme of the Cul de Sac Plain in Haiti. Two specimens from Port-au-Prince (MCZ 60154, MCZ 37599), one from Petionville (MCZ 60162) in the mountains south of Port-au-Prince, and three from Damien in the Cul de Sac north of Port-au-Prince (MCZ 60158-60) have a longitudinal pale lateral line (presumbly blue in life) on the lower- most scales rows ( scale row 2 in all, ex- panding to scale row 1 in one snake and to row 3 in two snakes ) . Other snakes from this general region ( Mont Caba'io, Morne Decayette, Morne I'Hopital, Delmas, Pen- eau — a total of 15 additional specimens) at present lack any indication of the lateral line. The snake from Peneau was examined while living and I can verify the absence of the lateral line. Thus, snakes from the area included by Morne Decayette in the west, Damien in the north, Delmas in the east, and Peneau and Mont Cabaio in the south I regard as intermediate in pattern between catesbyi and bariolatus. Snakes from the southern versant of the Massif de la Selle in this region ( Bascap-Rouge ) are typical U. c. catesbyi and show no bariolatus in- fluence. The Hispaniolan south island is comprised of both the Tiburon Peninsula in Haiti and the Peninsula de Barahona in the Repiiblica Dominicana. The north island includes the balance of Hispaniola north of the Cul de Sac- Valle de Neiba plain. In a large number of species of amphibians and reptiles, the south island area is occupied by populations subspecifically distinct from those of the north island. A reasonable assumption would be that U. c. catesbyi is the form occurring on the Peninsula de Barahona, but such does not seem to be the case. A series of 12 snakes from various Barahona localities, including the city of Barahona itself and its immediate environs, as well as localities (Oviedo, Peder- nales) well south on the peninsula, is avail- able; none of these snakes has the distinctive JJ. c. catesbyi lateral line. Two of the speci- mens were collected by myself and parties, and neither was recorded as having a pale lateral line; the balance of the Barahona ma- terial is from other collections and color and pattern data are not available. Thus I inter- pret the absence of the lateral line in all specimens from this region as a constant Barahona feature. The means of ventral scales in the Barahona snakes ( 164.5 in males, 164.4 in females) are quite compa- rable to similar means ( 164.8 and 164.4, respectively ) in catesbyi. There are no Bara- hona males with complete tails, but the fe- male subcaudal mean (179.0) is similar to that in female catesbyi ( 182.1 ). The largest female JJ . catesbyi examined (MCZ 'blK^^ ), with a snout-vent length of 908 mm, is from the Peninsula de Barahona; V. c. catesbyi is a large snake, although the largest Tiburon female has a snout-vent length of 830 mm. The Cul de Sac-Valle de Neiba plain pres- ently connects the previous Hispaniolan north and south islands. If this plain (in places lower than sea level ) was at times a strait separating the two islands and was recolonized after its emergence, it seems reasonable that the V . catesbyi populations in the plain would show a mixture of char- acteristics of the subspecies to the north and the south. I have pointed out above that bariolatus and catesbyi intergrade in the ex- treme western portion of the Cul de Sac. There is a series of 1 5 snakes from the balance of the plain ( localities from Eaux Gaillees in Haiti to Cabral in the Republica Dominicana, five of which were collected by Richard Thomas ) ; none of these shows any trace of a lateral pale stripe, nor any tendencies toward the nominate subspecies. Nine males have a ventral mean of 166.5 — intermediate between the mean ventral count of catesbyi + Barahona males on one hand, and bariolatus on the other. Six females from this region have a ventral count mean of 166.8, again intermediate between the same two samples. The subcaudal mean ( 1 86.8 ) of six males, however, is much lower than that of ten catesbyi males (194.6), although the extremes overlap very broadly; four females from the Cul de Sac-Valle de Neiba have a subcaudal mean (182.8) al- most identical with that of 1 1 female catesbyi (182.1). Specimens of V. catesbyi from Haiti {havi- No. 4 Uromacer catesbyi 147 olatiis ) differ from those from the Repiib- lica Dominicana {pcwipineiis) in number of subcaudal scales, the Dominican snakes with much lower ranges and means. It is possible that pampineiis ( with lower num- ber of subcaudals) has influenced the Cul de Sac-Valle de Neiba snakes to some extent, thereby accounting for the low number of subcaudals in males from the Cul de Sac- Valle de Neiba. I interpret the Cul de Sac-Valle de Neiba snakes as intermediate between catesbyi ( whose influence is most clearly shown in the laterally striped pattern at the extreme western end of the Cul de Sac), hariolatiis (whose influence is most clearly shown in the basic dorsal color and absence of a longi- tudinal line in the greater portion of the sample), and pampineus which has contrib- uted toward the lower number of subcaudal scales. The Barahona population is peculiar. In color and pattern it agrees with pmnpinens north of the Valle de Neiba, rather than with catesbyi to the west; in ventrals, the Bara- hona snakes are like catesbyi rather than pampineus. and in subcaudals they are like hariolatns (and intergradient specimens from the Cul de Sac-Valle de Neiba). Although there might be justification in the future for considering the Peninsula de Barahona populations as a subspecies distinct from all others, it seems prudent at this time to re- gard them as extreme intergrades between catesbyi and pampineus. The above discussion is based upon the following suite of specimens, grouped ac- cording to the areas mentioned: I. Port-au-Prince and vicinity {catesbyi X hariolatns ) : Haiti. Dept. de I'Ouest, Port- au-Prince (MCZ 60149-50, MCZ 60152- 57, MCZ .^7599); Morne Decayette (MCZ 62698 ) ; Source Leclerc, Morne Decayette ( MCZ 65969 ) ; Delmas ( MCZ 65970-71 ) ; Damien (MCZ 61058-61); Source Baria- joux, Morne I'Hopital (not mapped) (MCZ 65972); Petionville (MCZ 60162); Furcy, Mont CabaVo (MCZ 45743); Peneau, 5000 feet (1525 meters) (ASFS XI 572). II. Cul de Sac-Valle de Neiba ( catesbyi X hariolatus X paynpineus ) : Haiti, Dept. de I'Ouest. 13.1 km E Croix des Bouquets (ASFS V8144, ASFS V8345, ASFS V8307); Eaux Gaillees (MCZ 60168-70); Thomazcau ( MCZ 1 2864-65 ) ; Tete Source, 1.4 mi. ( 2.2 km) NNE Thomazeau (ASFS V8197); be- tween Thomazeau and Glore (MCZ 60171); Manneville (MCZ 8747); La Source, E of Fond Parisien (ASFS V8134); Repilblica Dominicana. Independencia Prov., Duvergc (AMNH 40993-94); Barahona Prov., woods at Cabral (AMNH 52397). III. Peninsula de Barahona ( catesbyi X pampineus): Repilblica Dominicana, Bara- hona Prov., Barahona (MCZ 43804-05); Palomino Springs, nr. Barahona (not mapped) (AMNH 59750); Valle de Polo (AMNH 51426-27, AMNH 51422); Hermann's finca, nr. Paraiso (AMNH 51424); 3.5 mi. (5.6 km) from Paraiso (AMNH 51428); Pederiidles Prov.. 13.1 mi. (21.0 km) S\V Enriquillo (ASFS V4425); Oviedo (MCZ 57765-66); 19 km N Pedernales, 1000 feet (305 meters) (ASFSV2697). Horn (1969) proposed an hypothetical history of the genus Uromacer which he con- sidered to be composed of four species. Uromacer catesbyi he regarded as the most primitive of the species, since it is the least specialized morphologically (all other spe- cies are long-snouted, attenuate snakes) and has a dietary reperatory which is composed of arboreal animals. (Presumably arboreality is the original niche for the genus, from which situation the remaining species have become more terrestrial. ) Horn further sug- gested that an ancestral colubrid (perhaps an unspecialized Alsophis — type snake) gave rise to both a pvoto-catesbyi and a proto- frenatus (the latter one of the long-snouted taxa ) , which were, respectively, the Uro- ?)iaccy of the north and south Hispaniolan islands. Although this is a possibility, the taxonomic status of the long-snouted forms precludes any more definitive statement of the early history of the genus. Whatever the early division of the genus, surely there was a basic division into short- and long-snouted forms, but I remain unconvinced that proto- catesbyi is the north island analogue of south island proto-frejhitus. It seems likely to me that, early in the history of proto-catesbyi, this group of snakes invaded the south island from the north island, and that the presence of a lineate lateral pattern in the Tiburon- Vache-Cayemites subspecies represents one old line of evolution from proto-catesbyi and the non-lineate subspecies represent the 148 Tulane Studies in Zoology and Botany Vol. 16 other. Although it is customary to consider that loss of a character is the more advanced condition (and thus the Tiburon-Vache- Cayemites subspecies would represent the primitive forms since they possess a pale lateral line), the non-lineate subspecies oc- cupy a much greater area and the absence of a lateral line would seem in this case to be the primitive rather than the advanced condition. Whatever the details, it seems certain that U. catesbyi was early split into two stocks, corresponding to the old north and south islands of Hispaniola. One of these ( that on the south island ) had a pale lateral longitudinal line ( catesbyi ) and from this stock arose insulaevaccariini and cereolinea- tHS. Both subspecies retain the lineate lateral pattern but have diverged in the color of the lateral line ( from blue to pale green or white). The north island stock, separated from its relatives on the south island by the then submerged Cul de Sac-Valle de Neiba plain, lacked a lateral longitudinal pale line but maintained a more or less uniform dorsal color ( although the lower sides may be lighter in contrast to the more dorsal colora- tion ) . With the elevation of the interisland strait, the south island catesbyi and the north island bariolatiis and pampineus once more came into contact and intergraded with one another. Intergradation is evident in the western Cul de Sac, where some snakes retain the blue lateral line of catesbyi and some do not. In the rest of the Cul de Sac-Valle de Neiba plain, the snakes present a mixture of scu- tellar characteristics derived from catesbyi, hariolatHS, and pampineus. The most unusual situation is that on the Peninsula de Bara- hona, where the snakes seem to represent in- tergrades between catesbyi and pajnpineus; typical blue-lined catesbyi occur no closer to the Republica Dominicana in the region of the Peninsula de Barahona than Bascap- Rouge on the southern slope of the Massif de la Selle in Haiti. Although the Barahona situation is not clear, it is possible that north- ern pampineus and southern catesbyi were long in contact in the Barahona region ( this contact possibly of longer duration than the final, complete closure of the interisland strait ) with consequently greater oppor- tunity for genetic mixing. There remains the possibility that, as in the case of the subspecies of Dromicus parvijrons ( Thomas and Schwartz, 1965:69-70), the Peninsula de Barahona populations of U, catesbyi show influence from an as yet undiscovered Isla Beata population, if it in fact exists. The two north island subspecies hariolatus and pampineus present a puzzle. I cannot arrive at any reason for the two subspecies to meet at approximately the Dominico- Haitian border as they appear to do. Along this border there is a more or less continuous montane barrier ( Montagues du Trou d'Eau- Sierra de Neiba; Massif du Nord-Cordillera Central ) ; these ranges lie at right angles to the international boundary and intermontane valleys should afford easy access between the two subspecies. Additionally, U. catesbyi occurs at high elevations, and mountains should not be effective barriers to these snakes. Two of the satellite subspecies, scandax and inchausteguii. differ from all other pop- ulations in having a patterned head ( ap- parently only in juvenile scandax). The islands inhabited by these subspecies, Tortue and Saona, lie diametrically opposed to His- paniola proper, one at the northwest (Tor- tue ) , the other at the southeast ( Saona ) . That such a pigmental trait occurs in two widely separated populations of U. catesbyi is perplexing. A comparable situation is the basically black ground color of all the satellite populations of Dromicus parvijrons, in contrast to most of the mainland snakes. Literature Cited Cochran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S. Natl. Mus. IHii- vii, 1-398, 120 figs., 12 pis. DOWLING, HerndoN G. 1951. A proposed standard system of counting ventrals in snakes. British Jour. Herpetology 1(5) :97-99, 1 fig- DuMERiL, A. M. C, AND Gabriel Bibron. 1854. Erpetologie generale, 7(l):i-xvi, 1- ""80, Paris. Dunn, Emmett R. 1920. On the Haitian snakes of the genera Leimadophis and Lho- niacer. Proc. New England Zool. Club "'i.^^— 44. Horn, Henry S. 1969. Polymorphism and evolution of the Hispaniolan snake genus Uro- macer (Colubridae) . Breviora, Mus. Comp. Zool. 324:1-23, 2 figs. Maerz, a., and M. Rea Paul. 1950. A dic- tionary of color. New York, McGraw-Hill Book Co., pp. i-vii, 1-23, 137-208, 56 pis. Mertens, Robert. 1939. Herpetologische Ergebnisse einer Reise nach der Insel His- No. 4 Urnmacer catesbyi 149 paniola, Westindien. Abh. senckcnberg. naturf. THOMAS, RICHARD. AND ALBERT SCHWARTZ. Ges. 449:1-84, 10 pis. I965. Hispaniolan snakes of the genus Drom- SCHLEGEL, HERMANN. 183 . Essai sur la phy- /f//j (Coiubridac). Rev. Biol. Trop. H (1 ) : sionomie des serpents. The Ha^ue, 2 vol., pp. co oi im r /-n/ .1 T 1 1 !^ ■> y ff 5J-,_J53 10 figs d()6, atlas, 21 pis. " SCHWARTZ, Albert. 1966. The Leiocephalus Williams, Ernest E. 1961. Notes on His- (Lacertilia, I^yuanidae) of Hispaniola. I. Leio- paniolan herpetolosjy. 3. The evolution and cephalus melanochlorus Cope. Jour. Ohio Her- relationships of the Anolis semilhieatus .qroup. petological Soc. 5(2):39-48, 1 fig. Breviora, Mus. Comp. Zool. 136:1-8. s-rufV-(\jL^ ^ \. TULANE STUDIES IN ZOOLOGY AND BOTANY VOLUME 16 INDEX TO AUTHORS AND SCIENTIFIC NAMES (New taxonomic entities in boldface) Acacia cym]}ispina, 91-92 Achatina fulica, 114 Acipenser fulvescens, 116 Agave falcata, 92 lechuguilla, 92 Alepisauriis brevirosiris, 117 Algansea monticola, 117 AhopJm, 147 AmblopUtcs ariommus, 126 Ameiva taeniura aequorea, 136 Anolis chlorocyatms, 133 coelestinus, 133 pccuariiis, 136 cybotes, 133 distichus, 133 fuhae, 136 Antigonia combafia, 125 Astatoreochromis aUuaudi, 19 Atriplex-Allenrolfea occidentalis, 91-92 Bascanion flagelliforme, 36 testaceum, 63 flagelliformis, 43, 51 fhgellum, 43, 63 frenatum, 33, 51, 58 laterale, 63 fidiginosum, 63 piceum, 51 flagelliforme, 36 bicinctum, 43 flagelliforme, 36 piceum, 51 iestaceum, 36, 51 testaceus, 47 flagellum, 51, 63 frenatum, 52 piceum, 52 Bcllaior ribeiroi, 128 Bradybaena similaris, 101-114 Brevoortia gunteri, 117 Bulimulus Cornells; 104 Bursera, 92 Calamus nodosus, 126 Carya spp., 42 Chilina cf fluminea, 103 Chirosioma riojai, 125 Cochlicella barbara, 104 Coluber constrictor mormon, 34 oaxaca, 33 siejnegerianus, 33 dipsas, 34 dorri, 34 flagelliformis testaceus, 36, 43 {f v^C flagellum, 33, 36 flagellum, 36, 43 fkwigularis, 45-46 frenatum, 63 k^ '^j frenatus, 51, 55 piceui', 35, 43, 51, 55, 58, 63 I '• w"%V.'A. ..ortcnburgeri, 33 siriolatus, 35 testaceus, 33, 36, 42 Corypliodon testaceus, 43 Cottus pygmaeus, 128 Crepis sp., 110 Crotalus viridis cerberus, 54 Cualac tessellattts, 125 Dendrophis catesbyi, 134 Dermatodidymocystis viviparoides, 27 Dcroccras laeve, 113 Diploglossus cosiatus nesobous, 136 stenurus, 136, 145 Draconetta oregona, 128 parvifrons, 145, 148 rosamondae, 136 Dryadophis, 34 Drymobius, 34 Dundee, Dee Saunders, article, 101-115 Elaphe frenata, 53 Eleutherodactylus pictissimus, 136 weinlandi, 145 Etheostoma coUettei, 126 ditrema, 126 microlepidum, 126 moorei, 126 nuchale, 127 radiosum, 127 nibrum, 127 Flourensia cernera, 92 Franseria dumosa, 92 catenatus, 1-21 grandis, 22-25 notatus, 16 stellifer, 1-21 Go/;/o.so?73a evelynac, 128 genie, 128 G(//£'//« /;/fo/o;-, 101-114 Gyrodactylus megacanthus, 22 prolongis, 22 trematoclithrus, 22 liNDEX TO AUTHORS AND SCIENTIFIC NAMES— Continued Hawaiia miniscula, 113 Hechtia sp., 92 Helicella, 104 Heliciiui orbiculata, 113 Hcrpetodnjas flagelliformis, 36, 42 flavigularis , 36 Heterodon platijrhinos, 89 Hijla dominicensis, 133 pulchrUineata, 133 Ictalurus serracanthus, 121 Jatropha, 92 Kamegai, Shunya, article, 26-29 Lame//flx/s gracilis, 101-114 mauritianus, 104 tnicra, 109 Lampropeltis getulus californiae, 75 triangulum, 54 zonata, 54 Lapoiiea canadcnsc, 110 Larrea tridentata, 91, 92 melanochlorus, 136 inclunocldorus, 133 Lepidodidymocystis irwini, 26—29 Lilium canadcme. 111 Limax marginatiis, 104 Liquidamhar siyracifhia, 42 Masticophis, 31-99 anthomji, 34, 35 aiiriguliis, 34 harbouri, 34 bilineaius, 34 flagelliformis, 36 tes-tacea, 43 testaceus, 36, 43 flagellum, 31-99 cingulum, 33, 35, 50-51, 58, 63, 72, 77, 87, 93-94 flagellum, 33-94 flavigularis, 33, 36, 43, 47, 55 frenatum, 33, 50-51, 53, 58, 63 frenatus, 51, 55, 58, 63 fuliginosus, 63-77, 80, 87, 92-94 Uneatulus, 33, 47-51, 56, 59-63, 7:3-77, 87, 91-94 picCTi-s, 31, 33, 35, 50-59, 64-77, 82, 87-94 ruddocki, 33, 35, 48, 53-58, 74, 87-94 testaceus, 34, 35-51, 59-73, 80, 87, 91, 93- 94 variolosus, 35 lateralis, 34 lim-atus, 34, 35 mentovarius, 34, 35, 93 picew.?, 51, 53-54, 58, 63 rutlweni, 34 schotti, 34 similineatus, 34 striolatus, 34, 35, 48 guardi, 34 ruthveni, 34 schotti, 34 taeniatus, 34 tesiaceus, 43 Menticirrhus nasus, 26—27 perigraptus, 113 thijroides, 114 A//crof/c577U/.s- corn, 128 suttkusi, 128 Myrica ccrifera, 42 Nocomis effusus, 117 asperifrons, 117 bailey i, 118 edwardraueyi, 118 euryzonus, 121 hypsdepis, 121 imeldae, 121 signipinnis, 121 uranoscopus, 121 albater, 122 elegans, 122 lachneri, 122 munitus, 122 phaeus, 125 stigmosus, 125 Ofiw o.s/0, 75 Oxybelis, 133 Panicum, 110 brachyanthus. 111 Fercino aurolincata, 127 leniicula, 127 Peristedion greyae, 128 Pffcea r?je.r/cflm/s, 125 echinata, 42 palusiris, 42 taeda, 42 Poblana fcrdebueni, 125 Poecilichthys whipplii radiosus, 127 Polyclithrum mugilini, 22-25 Po/!/«!/ra septemvolva, 113 texasiaiui, 113 Porichihys bathoiketcs, 128 Praticolella griseola, 113 Prietella phreatophila, 125 P/osop/s juliflora, 42. 92 flagelliform is, 36 Psammophis flavigularis, 33, 42, 45 Pupoides mmginatus, 113 Qwerctw falcata, 42 marilandica, 42 stellata, 42 INDEX TO AUTHORS AND SCIENTIFIC NAMES— Continued Rdchycentron canadus, 116 Retinella indentata, 113 Rogers, Wilmer A., article, 22-25 Rttmina dccollata, 101-114 Schwartz, Albert, article, 131-149 SnlaropsLs ruonile, 104 S})hoeruides parvus, 128 Starks-Ui y-lincata, 127 Strobdops texasiuna, 113 Succinea, 104 Suttkus, Royal D., article, 116-130 Stvingleus polyclithroides, 22—25 Thamnophis siifalis sirfalis, 89 Thcha piscina, 104 Thomerson, Jamie E., article, 1-21 Uromacer catesbi/i, 131-149 cateshi/i, 131, 134, 135, 137, 140, 144-148 cereolineatus, 131, 137, 138, 140, 144- 145, 148 frotiflicolor, 131, 137, 140, 142, 144-145 hariolatus, 131, 137, 138-139, 140-148 inchausteguii. 131, 137, 140, 143-145, 148 insulaevaccarum, 131, 136-137, 140, 144- 145, 148 pampineus, 131, 137, 138-141, 144-145, 147-148 scandax, 131, 137, 140-144, 148 dorsal is, 131, 133 fremitus, 131, 133, 147 oxi/rhi/nchm; 131, 133-134 scandax, 131, 134, 143 wetrtiorei, 131 Wilson, Larry David, article, 31-99 Yamaguti, Satyu, article, 26-29 Yucca, 92 Zamenis ihificUiforme, 36, 43 flaiicUiformis, 36, 43, 51 frenatus, 51 piceus, 51 flagcllum flagellum, 36, 43, 51 piceus, 51 flavigularis, 43 laterale fuliginosum, 63 lateralis fuUgimsus, 63-5, 70 Zonitoides arboreus, 104 ' I illllllllllllli 3 2044 093 361 053 Date Due I