Geographic Variation in the Yellow Mud Turtle, Kinosternon Flavescens

Home , Kinosternidae, Kinosternon, Yellow mud turtle

SHUTE ET AL.-WACCAMAW DARTER BIOLOGY 567

slenderhead darter, Percina phoxocephala, darters of the ingenus the Percina (Percidae) in the Kas- Embarras River, Illinois. Ibid. 74. kaskia River, Illinois. Ill. Nat. Hist. Surv. Biol. Note PFLIEGER, W. L. 1978. Distribution, 70. status, and life history of the Nianguae darter, VAN Etheostoma DEN ASSEM, J. 1967. nian- Territory in the three- guae. Missouri Dept. of Cons. Aquatic spined stickleback Series Gasterosteus 16. aculeatus L. Behavior RANEY, E. C., AND E. A. LACHNER. Suppl.1939. 16. Observations on the life history of the spotted darter, WANG, J. C.Poecilichthys S., AND R. J. KERNEHAN. 1979. Fishes of maculatus (Kirtland). Copeia 1939:157-165. the Delaware estuaries: a guide to the early life - , AND - . 1943. Age and growth of johnny histories. E. A. Communications. Townson, MD. darters, Boleosoma nigrum olmstedi (Storer) and Bo- WILLIAMS, T., AND B. C. BEDFORD. 1974. The use of leosoma longimanum (Jordan). Amer. Midl. Nat. otoliths for age determination, p. 114-123. In: 29:229-238. Aging of fish. T. B. Bagenal (ed.). Urwin Bros., SCHMIDT, R. E. 1979. Distribution and habitat of Surrey,the Eng. swamp darter (Etheostoma fusiforme) in southern WINN, H. E. 1958. Comparative reproductive behav- New England. Ibid. 408-413. ior and ecology of fourteen species of darters SPEARE, E. P. 1960. Growth of the central johnny (Pisces-Percidae). Ecol. Monogr. 28:155-191. darter, Etheostoma nigrum nigrum (Rafinesque) in Augusta Creek, Michigan. Copeia 1960:241-243. DEPARTMENT OF BIOLOGY, UNIVERSITY OF STARNES, W. C. 1977. The ecology and life history of NORTH CAROLINA AT WILMINGTON, WILMING- the endangered snail darter, Percina (Imostoma) tan- asi Etnier. Tenn. Wildlife Resources Agency. Fish- TON, NORTH CAROLINA 28406. PRESENT AD- eries Research Report. TWRA Technical Report DRESS (PWS AND JRS): DEPARTMENT OF ZOOL- 77-52. OGY, UNIVERSITY OF TENNESSEE, KNOXVILLE, THOMAS, D. L. 1970. An ecological study of four TN 37916. Accepted 28 May 1981.

Copeia, 1982(3), pp. 567-580

Geographic Variation in the Yellow Mud Turtle, Kinosternonflavescens

TIMOTHY W. HOUSEAL, JOHN W. BICKHAM AND MARLIN D. SPRINGER

Geographic and non-geographic morphological variation of 18 mensural characters of the shell, and color, is analyzed for populations of Kinosternonflavescens from throughout the range. K. f. arizonense and K. f. durangoense are found to be morphologically divergent from K. f.flavescens and K. f. spooneri by all statistical tests employed. K. f. spooneri is found to be indistinguishable from northern populations of K. f. flavescens and is placed in synonymy. K. f. durangoense and K.f. arizonense are slightly divergent from each other. The zoogeographic history of K. flavescens is discussed.

region. Kinosternon flavescens durangoense (Iver- K INOSTERNONflavescens wide-ranging turtle species is a whichcommon inhabits and son, 1979b) is known from the lower Rio Nazas the central and southwestern United States, region of Durango, and Coahuila Mexico; and and northern Mexico. Seven isolated popula- has also been reported from the Bolson de tions of K. flavescens, representing four subspe- Mapimi in the Mexican states of Durango, Chi- cies were recognized in the recent taxonomic huahua and Coahuila. Kinosternon flavescens revision by Iverson (1979b). Kinosternon flaves- spooneri (Smith, 1951) is restricted in distribu- cens arizonense (Gilmore, 1922; Iverson, 1979a) tion to relic sand prairies of Illinois, Iowa and is known from the drainages of the Rio Mag- Missouri. The nominate subspecies, K.f. flaves- dalena, Rio Yaqui, Rio Matape, Rio Sonora and cens (Agassiz, 1857; Hartweg, 1938) ranges the San Simon Valley, of the Sonoran desert from northern Nebraska south to northeastern 568 COPEIA, 1982, NO. 3

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Mexico (Iverson and length,Berry, M9L; marginal 1979), nine width,and M9W; west max- to Colorado and southeastern Arizona. The dis- imum plastron length, PL; plastron width at tribution of K. flavescens and other Kinosternon humeral-pectoral seam, PW1; plastron width at species in the southwest is discussed in detail pectoral-abdominal by seam, PW2; plastron width Iverson (1978) and Conant and Berry (1978). at abdominal-femoral seam, PW3; gular length, The presence of K. flavescens in Illinois GL; or gular width, GW; inter-humeral seam Iowa was first reported by Cahn (1931, 1937), length, IHSL; inter-abdominal seam length, at which time he noted its darker coloration. IASL; inter-femoral seam length, IFSL; inter- These populations received formal subspecific anal seam length, IANSL; forelobe length, recognition (K. f. spooneri) in 1951 (Smith, FLL; hindlobe length, HLL. These characters 1951). The Illinois Mud Turtle, K. f spooneri, are those which were found to be most useful was recently proposed for addition to the fed-in Kinosternon taxonomy by Iverson (1979b) eral Endangered Species List (Federal Register, and Berry (1978). Only specimens greater than 6 July 1978; Brown and Moll, 1978). A clear 80 mm CL, and possessing the complete com- understanding of the systematic status of en-plement of characters, were included in the sta- dangered, and proposed endangered species tistical is analyses. Specimens larger than this essential before appropriate management couldde- be easily sexed on the basis of secondary cisions can be made. The purpose of this study sex characteristics and were assumed to be sex- is to analyze the extent of geographic variation ually mature. All specimens were measured by in K. flavescens, specifically to clarify the taxo- TWH with the exception of 140 specimens of nomic status of K. f. spooneri. K. f. spooneri (including all 24 Illinois River specimens) which were mark-recapture speci- MATERIALS AND METHODS mens, and measured by employees of LGL Eco- logical Research Associates, Inc., who were A total of 961 specimens of K.flavescens, from studying those populations. Sixteen character throughout the geographic range of the species ratios were computed (carapacial measure- (Fig. 1), was examined, and eighteen characters ments being expressed as a ratio of CL, and of the shell measured with dial calipers to plastralthe measurements being expressed as a ra- nearest 0.05 mm. These characters and their tio of PL), to account for differences in absolute abbreviations are as follows: carapace length, measurements resulting from indeterminate CL; carapace width, CW; vertebral one length, growth. V1L; vertebral one width, ViW; marginal nine For ease in the statistical analyses, specimens HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 569

TABLE 1. SPECIMENS USED IN THE STATISTICAL ANALYSES, ARRANGED INTO SAMPLES. Sample localities are mapped in Fig. 1.

Sample size Sample Males females Total

K. f spooneri 1. Illinois R. 14 10 24 2. Mississippi R. 120 101 221

K. f. flavescens 3. Chariton R. 1 0 1 4. Nebraska Sandhills 8 12 20 5. Blue R. 0 1 1 6. Republican R. 9 15 24 7. Solomon R. 0 3 3 8. Smoky Hill R. 17 17 34 9. Arkansas R. 14 46 60 10. Neosho R. 3 0 3 11. des Cygnes R. 1 1 2 12. Salt Fork of Arkansas R. 27 50 77 13. Cimarron R. 27 27 54 14. Canadian R. 55 76 131 15. Red R. 27 19 46 16. Beaver R. 28 24 52 17. Upper Brazos R. 18 16 34 18. Lower Brazos R. 2 5 7 19. Upper Colorado R. 5 1 6 20. Lower Colorado R. 1 1 2 21. Lavaca R. 0 1 1 22. Guadalupe R. and San Antonio R. 3 1 4 23. Blanco-Medio Creek 0 1 1 24. Nueces R. 1 2 3 25. Pecos R. 27 27 54 26. Upper Rio Grande 10 3 13 27. Lower Rio Grande 2 2 4 28. Playa Lakes 12 8 20 29. Whitewater Creek 1 0 1 30. Rio Case Grandes 0 2 2 31. Rio Santa Maria 7 4 11 32. Rio Conchos 4 2 6 33. Rio Salado 11 12 23

K. f. arizonense 34. Rio Magdalena 1 5 6 35. Rio Yaqui 1 0 1 36. Rio Matape 1 1 2 37. Rio Sonora 1 4 5 38. San Simon Valley 1 4 5 K. f durangoense 39. Rio Nazas 3 2 5 40. Bolson de Mapimi 0 2 2

were arranged intherefore, samples (Fig.this arrangement1, Table 1) cor- is both a conve- responding to nientmajor anddrainage effective systems. means Kino- of grouping data sternon distribution (Berry, tends 1978; to be Iverson,limited by 1979b). major These samples river drainage function basins or as their Operational subdivisions; Taxonomic Units 570 COPEIA, 1982, NO. 3

(OTU's) in the statistical sisted of Olive-Brown, analyses. Brownish-Olive, It Oliveis not known if the specimen and Grayish-Olive from samplecolored specimens 3 repre-(light), sents a natural population. while Class II contained J. Olive-Green L. Christiansen to Green- (pers. comm.) informed ish-Olive usand Blackishthat Neutral this Gray may specimens be an introduced individual (intermediate); of K. and f.Class flavescens. III contained Dusky We have therefore included Brown, Darkit Grayishwith Brown this and Fuscous subspecies spec- in Table 1. imens (dark). Color names follow Smithe Programs from the Statistical Analysis System (1975). (SAS), designed and implemented by Barr and Goodnight (Service, 1972), and NT-SYS (Rohlf Data standardization.-Many arguments, both and Kishpaugh, 1972) packages were used pro in (Corruccini, 1977; Dodson, 1978; Hills, the statistical analyses. Multivariate Analysis 1978) of and con (Atchley et al., 1975, 1976; Atch- Variance (MANOVA) was performed using ley and Anderson, 1978; Albrecht, 1978) have SAS, whereas the Principal Components Anal- appeared in recent literature concerning the ysis (PCA) and cluster analysis employed use NT- of ratios in statistical analyses. Atchley et al. SYS. Univariate analyses using SAS included (1976) recommended the use of residual values the Student's t-Test, Analysis of Variance from(AN- a regression analysis (over ratios) as input OVA), and Duncan's Multiple Range Test. variables To when standardization is necessary. test for sexual dimorphism, the Student's t-Test The utility of both methods was discussed by was performed on data from a series of 96 spec-Atchley (1978). Berry (1978) tested these two imens of K. f. spooneri from the Big Sandmethods of standardization (ratios vs regres- Mound population (Muscatine-Louisa counties, sion) in multivariate analyses of other Kinoster- Iowa) for which voucher specimens are avail-non species, and found virtually no differences able. in the results obtained. The use of character All of these analyses are conservative tests ratios as input variables seems justified (Iver- requiring little or no a priori assumptions re- son, 1979b; Berry, 1978), and are used in this garding the data. MANOVA-canonical analysis study. assumes that specimens can be grouped into Acronyms follow Duellman, Fritts and Levi- samples. Mean sample values from the MAN- ton (1978) with the following exceptions: DU = OVA-canonical analysis for vectors one and two Drake University, JSA = John S. Applegarth, were plotted, and ellipses approximating one University of New Mexico, MHP = Museum of standard deviation were drawn around the the High Plains, Fort Hays State University, plotted means. The cluster analysis provides NWOSU a = Northwest Oklahoma State Univer- phenogram depicting the similarities amongsity, RS = Robert Semmler, University of New the sample means. Since different clustering Mexico, TTU = The Museum, Texas Tech routines result in different phenograms, University, the TWH = Timothy William Houseal, UPGMA cluster routine on standardized dis- field series, Texas A&M University, UAZ = tance coefficients was used. This is probably University the of Arizona, Department of Zoology, most appropriate clustering routine available UF-FSM = University of Florida, Florida State (Sneath and Sokal, 1973). Principal Compo- Museum and UOMZ = University of Okla- nents Analysis is very conservative and provides homa Museum of Zoology. an accurate representation of the distances between major groups (Rohlf, 1968). Specimens examined.---Specimens marked with an asterisk were exam- Geographic variation in color was analyzed ined, but were not included in the statistical analyses. because of its historic importance in the rec-Kinosternonflavescens spooneri: Illinois River. Illinois. Mason Co., Sni- ognition of K. f. spooneri. Color is difficult carte to Slough, MCZ 53932* (paratype), FMNH 37992* (paratype); 5 km NE Havana, UMMZ 103089* (paratype), INHS 6010-11* (paratypes). quantify due to the qualitative nature of Mason the and Tazewell Co's., 24 mark-recapture specimens, LGL Ecolog- character and inconsistencies in preservation ical Research Associates. Cass Co., 1.6 km S Beardstown, INHS 5987- techniques. Only those specimens collected 89* (paratypes). Morgan Co., Meredosia Bay, UMMZ 47654* (paratype), INHS 5587-89*, UIMNH 2252-53*, 2255-56* (paratypes), during the summer of 1979, a total of 102 USNMspec- 83190* (paratype). Henderson Co., 11 km N Oquawka, INHS imens grouped into seven localities (Fig. 4244* 11), (holotype), 4245* (paratype), 3220-22* (paratypes). and preserved under identical conditions, Mississippiwere River. Iowa. Muscatine-Louisa Co's., Big Sand Mound, 96 included in the color analysis. The specimens specimens, DU unnumbered, 90 mark-recapture specimens, LGL Eco- were arranged from light to dark, and logicalthen Research Associates. Missouri. Clark Co., 35 mark-recapture specimens, LGL Ecological divided into three classes (Fig. 2). Class IResearch con- Associates. HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 571

Fig. 2. Representatives of the three color classes used in the color analysis. Class I (top; left to right, TWH 169, 293, 310), Olive-Brown to Olive and Grayish-Olive colored specimens; Class II (middle; left to right, TWH 166, 168, 154), Olive-Green to Greenish-Olive and Blackish Neutral Gray colored specimens; Class III (bottom; left to right, TWH 173, 134, 122), Dusky Brown to Fuscous colored specimens. Color names follow Smithe (1975). 572 COPEIA, 1982, NO. 3

Chariton River. Missouri. Adair Oklahoma. Co., Major16 Co.,km KU 18382;SW 5 kmKirksville, S Cleo Springs, UOMZ DU unnumbered. 8611-12; 11 km E Orienta, UOMZ 8596. Woods Co., UOMZ 8538-39; 3 km W Edith, UOMZ 9510, 9614-17, 9628, 9630, 9673-74, 9683-84, Kinosternonflavescensflavescens: Nebraska Sandhills. Nebraska. Cherry 9707; 4.8 km W Edith, UOMZ 9703-04; 3 km NW Edith, UOMZ 9658; Co., Three Cornered Lake, 48 km N Hyannis, UN 6774; Valentine, 3 km SW Waynoka, UOMZ 8537. Kingfisher Co., 10 km E Kingfisher, UN 1982; Valentine National Wildlife Refuge, Pelican Lake, TCWC UOMZ 5389. Woodward Co., 11 km SW Freedom, UOMZ 9681. Ci- 58183-58256, 58258-58263, 58265, 58274, 58281, 58291, 58300, marron Co., 29 km E Kenton, UOMZ 5294-96, 5298; 5 km N Kenton, 58308. Garden Co., 45 km N Oshkosh, UN 7861-62. UOMZ 5136.

Blue River. Nebraska. Kearney Co., 5 km NE Minden, UMMZ Canadian River. Oklahoma. Dewey Co., 8 km SW Canton, KU 18128; 101295*. 6 km SW Taloga, UOMZ 8473. Cleveland Co., Norman, UOMZ 1472, 3755, 5876, 6422, 6455-56; 1.6 km S Norman, UOMZ 19913-14, Republican River. Nebraska. Furnas Co., 3 km SE Cambridge, 22860; UN 3 km S Norman, UOMZ 467, 19912, 15265; 8 km SW Norman, 1984-85. Harlan Co., 11 km E Oxford, UN 1558. Hitchcock Co., UN UOMZ 468; 11.3 km E, 3 km S Norman, UOMZ 20223; 2.4 km NW 1560. Dundy Co., Rock Creek State Fish Hatchery, 11 km N Parks, UN Norman, UOMZ 19022; 3 km N Norman, UOMZ 2714; Indian 1652-57, TCWC 58235, 58236, 58248-58250, 58252, 58280, 58287, Springs, UOMZ 19094; 0.8 km N Midway, UOMZ 19097. Oklahoma 58295, 58305; Rock Creek State Recreation Area, 10 km N Parks, Co., 1.6 km W, 2.4 km S Edmond, UOMZ 20219. McClain Co., UOMZ TCWC 58251, 58272. 19048-49. Ellis Co., 11 km E Arnett, UOMZ 24281-83. Kansas. Cheyenne Co., 20 km W, 13 km S St. Francis, KU 174648- Texas. Hartley Co., 48 km SW Dalhart, KU 72809. 49. New Mexico. Quay Co., 0.8 km SE Ute Creek Dam, UNM 13283- 86, 13290-99, 14190, 20690, 20694, 20700-01, 22388, 22392, 23180- Solomon River. Kansas. Ottawa Co., State Lake, KU 174764. Graham 81, 31949-52, 31954-58, 31968; 10 km S, 6.8 km W Logan, UNM Co., 10 km E Hill City, MPH 297. Phillips Co., 6 km S Glade, KU 20511, 20517-19, 20525, 20685, 20688, 20691, 20695, 31959-67; 19480. North Shore Lake, E Tucumcari, UNM 20512, 20516, 20530, 20554, 20575, 20577-78, 20579 (2), 20580-86, 20588-89; 1.6 km below Ute Smoky Hill River. Kansas. Saline Co., 14 km S Brookville, KU 174669. Creek Dam, UNM 17192. Union Co., 34 km N Nara Vista, UNM Logan Co., 10 km S Elkader, KU 3753-54, 3761. Wallace Co., 23747-49, KU 31969; 11 km S Clayton, UNM 20692, 20696-97, 20699, 3021-23. Russel Co., 3 km W Dorrance, MHP 65. Trego Co., R22W, 23189, 23750-51; Capulin National Monument, UNM 23169. San T15S, Sec 1, MHP 5341-42; East Side Cedar Bluff Dam, MHP 4561. Miguel Co., near Conchos Dam State Park, UNM 11257, 17192, 21167. Ellis Co., R19W, T14S, Sec 1, MHP 6116; 8 km W Hays, MHP 1438, Harding Co., 2.5 km due W Ute Creek bridge on NM 39, 19 specimens, 1440, 1472; 18 km S Hays, MHP 2724; 39 km W, 22 km S Hays, MHP RS unnumbered. 1439; 3 km W, 14 km S Hays, MHP 2384; 11 km S, 16 km W Hays, MHP 300; 6 km S, 4 km W Antonino, MHP 3594-95; 5 km S, 3 km Red River. Oklahoma. Greer Co., between Mangum and Reed, UOMZ W. Antonino, MHP 4870, TCWC 58241-43, 58246, 58293, 10 km W 28625; 8 km Wjct hwy 9-238, UOMZ 23809, 23821, 23830, 23833; 1.3 Hays, TCWC 58238-58240, 58268, 58271, 58283, 58317; 1 km NW km N Reed, UOMZ 23875-76; 1.6 km E Plainview, UOMZ 23870, Schoenchen, TCWC 58244. 23872. Cotton Co., Walters, UOMZ 23765. Garvin Co., Mayesville, UOMZ 10935; 1.6 km W Pooli, UOMZ 23798. Beckham Co., 3 km S Arkansas River. Kansas. Rice Co., 5 km S Sterling, MHP unnumbered; Carter, UOMZ 13046, 13049-50, 13055, 13060-61, 13063, 13066, 14 km N, 6 km W Hutchinson, KU 154064. Reno Co., 2 km N Pleuma, 13068, 13070-74, 13076; 8 km S Carter, UOMZ 13039; 7.2 km SE MHP 4556; 11 km S Sterling, MHP 1416; Hutchinson, KU 153041. Erick, UOMZ 23880. Roger Mills Co., UOMZ 4702. Tillman Co., 8 km Barton Co., 16 km N Great Bend, MHP 4559. Stafford Co., R11W, E Davidson, UOMZ 12999. Kiowa Co., Kiowa/Till Lin, UOMZ 8423- T22S, SE ? Sec 19, MHP F43; 13 km N, 10 km E Stafford, KU 41722- 24. Jefferson Co., 1.4 km E jct hwy 70-81, UOMZ 25284. Comanche 24; 19 km N, 10 km E Stafford, KU 41734, 69986-95, 69998-70003, Co., 13 km SW Lawton, UOMZ 19995, 19999; 18 km SW Lawton, 70005-10; 24 km N, 10 km E Stafford, KU 41725-27, 41729; 16 km UOMZ 19989. Harmon Co., 6 km E Hollis, UOMZ 4616; 13 km SE N, 8 km E Fowler, KU 50304, 50306-07, 50309; Little Salt Marsh, KU Hollis, UOMZ 4594-95, 4602, 4604, 4607, 4611-12; 3 km E Vinson, 3105-06, 3165-67, 3169-72, 3255. Kingman Co., Kingman, KU 18386, UOMZ 23820. 18388; 1 km S Runny Meade, KU 17212. Cowley Co., 3 km S, 1 km Arkansas. Red River, MCZ 1919* (cotype). W Udall, KU 3171, 51450. Edwards Co., 5 km S Troosdale, KU 3032. Ford Co., 2 km ENE Bloom, UNM 13277; 5 km ENE Bloom, UNM Beaver River. Oklahoma. Texas Co., 6 km S Hooker, KU 46869. Bea- 13278; 6 km ENE Bloom, UNM 13279. ver Co., 20 km W Knowles, TCWC 58282, 58294; 4.5 km E Gate, UOMZ 5426, 24222, 24234, 24242, 24245, 24247, 24249, 24251-52, Neosho River. Missouri. Barry Co., 5 km W Exeter, KU 177109. 24254-69, 24271-72, 24274-75; 3.2 km E Gate, UOMZ 5412, 5414- Kansas. Cherokee Co., 6 km N Riverton, KU 23356. Greenwood Co., 15; West-central part of county, UOMZ 25028. Cimarron Co., 11 km MCZ 8432*. S Boise City, UOMZ 5010, 5123, 5028, 5056, 5075, 5111, 5135. Harper Oklahoma. Mayes Co., 3 km E Chouteau, UOMZ unnumbered. Co., UOMZ 13752; 1.6 km E Rosston, UOMZ 24216. Woodward Co., 8 km NE Woodward, UOMZ 8570, 8572-73, 8584, 8586, 8588. Des Cygnes River. Kansas. Franklin Co., KU 3029-30. Upper Brazos River (above Palo Pinto Creek). Texas. Lubbock Co., Salt Fork. Kansas. Barber Co., 2 km W Aetna, KU 20520-21; Lubbock, 10 km TTU 464-1 to 464-4, 960-1 to 960-6, TCWC 58289, 58292, N, 6 km E Sharon, KU 41712; 10 km N Hartner, NWOSU unnum- TWH 284. Crosby Co., about 11 km S Lorenzo, TCWC 58206-58217, bered. Kiowa Co., 14 km SW Belvidere, KU 18379. 58279, 58286. Floyd Co., 6 km S Floydada, TTU G-5812. Eastland Co., Oklahoma. Alfalfa Co., 5 km E Cherokee, UOMZ 9135-36, Cisco,9138- KU 3130. Callahan Co., 5 km NW Baird, TCWC 21406. Young 42, 9171, 9179, 9187-93, 9195, 9198-200, 9203, 19067-71, Co., 19073, 29 km N Graham, TCWC 562. Baylor Co., 13 km SSE Seymour, 19075, 19077-78, 19096; 11 km SE Cherokee, UOMZ 9300; TCWC 3 km 54107; SE 16 km W Seymour, TTU G-5801. Cherokee, UOMZ 9246; State Fishery Hatchery, UOMZ 19080-85; Great Salt Plain, UOMZ 25268, 25271-72, KU 18375, UOMZ Lower 5401, Brazos River (below Palo Pinto Creek). Texas. Erath Co., 1.6 5403; 2.4 km S Byron, NWOSU unnumbered; 14 km W, 5 km km S NCher- Stephenville, TCWC 13963; 0.8 km W Stephenville, TCWC okee, NWOSU unnumbered. Woods Co. Alva, Hatfield Lake, 20062. TCWC Hood Co., 4.0 km N Lipan, TCWC 25124; 3 km E Lipan, 58220, 58223-58226, 5 km N Alva, TCWC 58231; 6 km N, TCWC 2 km 25120,W 25122. Brazos Co., TCWC 58290; 3 km NE Bryan Alva, 3 specimens, NWOSU unnumbered; about 8 km NW courthouse, Alva, TCWC 44478. TCWC 58227-58230, 58232, 58273; 5.5 km Sjct US 281 and OK 11, TCWC 58310. Upper Colorado River (above Pedernales River). Texas. Lynn Co., 32 km S Tahoka, TTU G-5811. Menard Co., Menard, TCWC 42342. Cimarron River. Kansas. Morton Co., Elkhart, KU 3037, 3040-45, Midland Co., 20.1 km S Midland, TCWC 27546; 18.8 km S Midland, 3047-51. Clark Co., 2.4 km ENE Minneola, UNM 13281. Meade Co., TCWC 27547. Mason Co., 6 km SW Mason, TCWC 31273; 11 km SW 6 km W Fowler, KU 23081; 11 km N Englewood, KU 18392. Mason, TCWC 31270. HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 573

Lower Colorado River (below Pedernales River). Texas. Wharton 48534-543; Co.,3 km W Aguajita, FMNH 47362-63; Nueva Rosita, FMNH 5 km W El Campo, TCWC 18426-27. 47364.

Lavaca River. Texas. Jackson Co., 3 km W Granada, TCWC Kinosternon 14577. flavesces arizonense.--San Simon Valley. Arizona. Pima Co., W Sells, JSA unnumbered; 6.8 km E Sells, UAZ 27954; 3.5 km E Guadalupe-San Antonio rivers. Texas. Kerr Co., 24 km Sells, S Mountain UAZ 27949, 27955; 4 km E Sells, UAZ 27950. Home, TCWC 36611. Goliad Co., 13 km SW Goliad, TCWC 52118. Bexar Co., 16 km S San Antonio, TCWC 14950. Refugio Co., 13 km Rio Magdalena. Sonora. About 26 km S Santa Ana, UAZ 31739-40; N Tivoli, KU 44945. Hays Co., San Marcos, UMMZ 118271*. Rio Blan- Desenboque del Rio, UAZ 43037; Llano, UMMZ 72234 (K.f stejnegeri co, USNM 50* (lectotype), 131823* (lectoparatype). paratype), 72235 (K. f stejnegeri holotype); Trincheras AMNH 67502.

Blanco-Medio Creek. Texas. Bee Co., 11 km W Beeville, TCWC 52117. Rio Yaqui. Sonora. 13 km SE Tecoripa on rd to Tonichi, UAZ 17957.

Nueces River. Texas. La Salle Co., 3 km S Woodward, TCWC 333, Rio Matape. Sonora. 15.6 km W Tecoripa, UAZ 27958; 5 km E Ma- 459. Live Oaks Co., 9.2 km W Jim Wells Co. line, TCWC 21405. zatan, UNM 17663.

Pecos River. New Mexico. Chavez Co., Dexter National Fish Hatchery, Rio Sonora. Sonora. 26 km N Hermosillo, UIMNH 73495-97; 72.6 km TCWC 58301-58302, 58311-58312, 58315-58316, 58319; 4 km N S Santa Ana, UIMNH 23919-20. Twin Lakes, UNM 15564; Cockleburr Lake, 8 km W Pecos R, UNM 16138; 49.7 km E turnoff to Bottomless Lakes State Park on US 380, Gila River. Arizona. Gila River, Camp Yuma, USNM 7892* (K.flaves- UNM 14741-42; 0.8 km E of N entrance to Bottomless Lakes State cens lectoparatype). Park, UNM 23745; 6 km NW Bottomless Lakes State Park, UNM 12506; 1.2 km NW Bottomless Lakes State Park, UNM 12507-09; 0.4 Kinosternon flavescens durangoense.-Rio Nazas. Durango. 22 km NE km N Bottomless Lakes State Park, UNM 13013; Bottomless Lakes Pedricena, FMNH 112996-98, 123666, 179224 (paratypes), UIMNH State Park, UNM 12504, 15565-68; 16 km NE Hwy 70 Pecos R bridge, 19338 (paratype), 19339. UNM 14870; 16 km N rte 7 Pecos R bridge, UNM 14612-13; 16 km E Roswell, UNM 12503; Country Club Lake, NE Roswell, UNM 14489, Bolson de Mapimi. Durango. Ceballos, 8 km from Torreon, UF-FSM 14746; 1.2 km S bridge on rd to Dexter, UNM 12505; 10.3 km W 16180 (holotype). turnoff to Bitter Lakes State Park, UNM 14865; Bitter Lakes, NE Ros- Chihuahua. 10 km N Escalon on Hwy 49, CM 59376. well, UNM 15030, 15563, 17208-09, 23363, 30755-56. Eddy Co., 0.5 Coahuila. Jaral, UMMZ 117425*. km W Avalon Dam, UNM 23746; 16 km W Malaga, UNM 20485; 1.6 km below Harroun Dam, UNM 7520; Pecos R at NM 31 bridge, UNM K.f. durangoense x K.f.flavescens.-Rio Florido. Chihuahua. Las Are- 23173; R25E, T22S, W ? of SE ? of Sec 23, UNM 23175, 23177; nosa, Laga de Las Arenosas, KU 33914*. R25E, T21S, NW ? of Sec 17, UNM 23165-66. Guadalupe Co., Twin Lake, 1 km S, 2.3 km E US 66 Pecos R bridge, UNM 20608; NE shore, Perch Lake, 1.4 km S, 2.4 km E US 66 Pecos R bridge, UNM 20604- RESULTS AND DISCUSSION 05, 20686; Rock Lake 3.5 km S, 1.9 km E US 66 Pecos R bridge, UNM 20606-07; Rock Lake Fish Hatchery, 3 km S Santa Rosa, UNM 20535. Non-geographic variation.-Sexual dimorphism Texas. Pecos Co., Ft. Stockton area, KU 8860. Reeves Co., San Sol- omon Springs, Balmorhea State Park, UAZ 27948. in a series of 96 specimens of K.f spooneri from Big Sand Mound, Iowa was significant (P < .01) Upper Rio Grande (above Pecos River). New Mexico. Dona Ana Co., in 12 of 16 character ratios. The five most im- 0.4 km N Garfield exit of 1-25, UNM 20527, 20591-96, 20598. Texas. Brewster Co., Rio Grande Campground, Big Bend National portant characters in distinguishing the sexes Park, UNM 16695, 17191; Dagger Flats, Big Bend National Park, KU are (in descending order) IANSUPL, CW/CL, 177110; Rio Grande Village, Big Bend National Park, KU 177111. HLL/PL, and PW3/PL. The females have a rel- Presidio Co., 56 km SSE Marfa, UMMZ 10129-93*. Chihuahua. 26 km S Cd. Juarez, KU 45018. atively longer IANSL, and a relatively wider carapace (CW). Due to this sexual dimorphism, Lower Rio Grande (below Pecos River). Texas. Brooks Co., 20 km SW Falfurrias, TCWC 20907. Starr Co., 23 km E Rio Grande City, TCWC sexes were treated separately in all subsequent 18429. Cammeron Co., 3.9 km E FM 2480 on FM 510, TCWC 52375; analyses, except color. Port Isabel, TCWC 53103. Rio Grande, MCZ 1918* (cotype). Coefficients of variation (CV) serve as a good Playa Lakes. New Mexico. Hidalgo Co., Pond S NM 9, 18.0 km E jct measure of individual variation. Coefficients of with NM 338, UNM 20513-15, 20520-23, 20526, 20531-34, 20553, variation of the 16 character ratios used in this 20599-603; 26 km S Hachitas on NM 81, UNM 23754; Playa Lake 12 km S NM 9, 18 km E Animas, UNM 23753. study in a series of specimens from Sample 2 (Mississippi River, N = 120 d&d, 101 909) Whitewater Creek. Arizona. Chochise Co., about 35 km N Douglas, near Elfrida, UAZ 27953. ranged from 3.41 (FLL/PL) to 35.51 (IFSL/PL) for males; and from 2.85 (CW/CL) to 46.32 Rio Case Grandes. Chihuahua. 24 km S Ascension, UAZ 36375-76. (IFSUPL) for females. IFSL/PL had the highest Rio Santa Maria. Chihuahua. Near Progresso, UMMZ 118273-283. mean CV (from all samples) for both males and females. Other highly variable character ratios Rio Conchos. Chihuahua. 22.4 km E Aldama Pemex on rte 16, UNM 31946; 21.4 km E Aldama Pemez on rte 16, UNM 31948; 13.4 km E for both sexes include (in descending order) Aldama Pemex on rte 16, UNM 31943; 12.6 km E Aldama Pemex on GL/PL, IHSL/PL, M9W/CL, and GW/PL. The rte 16, UNM 31375; 2.9 km E Aldama Pemex on rte 16, UNM 31947; least variable character ratios (lowest mean CV El Rinco, 18 km W Ojinaga, UAZ 35030. values) for both sexes are (in ascending order) Rio Salado. Coahuila. 3 km N Nava, FMNH 28840; 5 km NW San HLL/PL, FLIPL, PW2/PL, and CW/CL. Males Juan, FMNH 47355-56; 5 km W Allende, FMNH 47357-59, 47365; 8 km S Sabinas, FMNH 47360; 5 km S Sabinas, FMNH 47361; 10.9 had higher mean CV's for seven character ra- km E Sabinas, UIMNH 62779-780; 22 km E Sacramento, UIMNH tios, and females for the remaining nine. 574 COPEIA, 1982, NO. 3

MALES FEMALES

815 2 120 2 101 101- 13 llt

12 ..... 27 12 . so 33 .25 MALES 13_ .n 27 13.6?. 27

25- 3 27 25 =in 26A .510 26A-O3 28 l 12 28 - 8 34-38 6 34-38 12 Q s22~ 0.35I I03 0.39 0.43I i 0470 ,0.51 I 0.55i , 0,390 , 0.43 I 0, 0.47 . 00S3. I 0.0I Fig. 3. Modified Dice-Leraas diagram for PW3/PL ?Wi 31 2 r \ along a northeast-southwest transect. Horizontal line represents range, vertical line represents mean, solid rectangle represents one -65 standard deviation and open rectangle represents two standard errors. Num- ber to left is sample number, while32 28 19ON:3.ff the ~0 1number to right indicates sample size.

Geographic variation-univariate -50 40 VECTOR statistics.-All ONE; 37.27% "20 10 16 character ratios in the ANOVAFig. 5. Two-dimensional procedure forplots of the mean of the males had significant F canonical values variates atfrom thethe MANOVA-canonical P = .001 level. For the females, allanalysis character for males from allratios samples. except M9L/CL had F values which were significant at the P = .001 level. These results indicate significant geographic variation inthe all range (samples 10-25), with smaller values character ratios except M9L/CL for females. being found in the northeast (1-11) and south- Modified Dice-Leraas diagrams for PW3/PL west samples (26-38). Variation in IANSL/PL (Fig. 3) and IANSL/PL (Fig. 4) along a north- (Fig. 4) shows a slight clinal decrease in size toward the southwest. The variational trends of east-southwest transect from the K. f. spooneri samples to the K. f. arizonense samples, are these pre- two character ratios (PW3/PL and IANSL/ sented. These character ratios (PW3/PL PL) and reflect the trends seen in the other char- IANSL/PL) are highly influential on vectors acter ratios. Little clinal variation exists; rather, one and two in the MANOVA-canonical anal- the means of the samples fall within a wide ysis and factors one and two in the PCA, rangere- of variation for each character ratio in spectively. For PW3/PL (Fig. 3) there appears the K. f. spooneri and K. f flavescens samples (1- to be an increase in size toward the center of 28). However, the mean of the combined K. f arizonense samples (34-38) falls outside one standard deviation of the means of all K. f.

MALES FEMALES spooneri and K. f. flavescens samples.

Geographic variation-multivariate statistics.-A 2 . .... 120 2 10 MANOVA-canonical analysis was performed 1213-i2 -." - 27 13il 12 A 27s o with all samples, for both males and females separately. Four different criteria (Hottelling- 16 W 20 s s 24 Lawley's Trace, Pillai's Trace, Wilk's Criterion 14 , 55 14 ._ ib 76 and Roy's Maximum Root Criteria) were used n26 . ~ 27lo0 25 261 -. ' 1328 to test the hypothesis of no overall locality ef- 28?-.L612 28,,. s . fect. F values from all four tests were significant 34. 6 -- 34 .3 ---44 12 at the P = .0001 level. Morphological variation 0.14 0.18 0.23 0.20 0.30 0.34 0.17 0.21 05 0.29 0.33 0.37 among samples, for both sexes, is therefore as- Fig. 4. Modified Dice-Leraas sumed diagram to exist due to forthe effect IANSL/ of locality. PL along a northeast-southwest The first transect. two canonical Explanation variates were com- as in Fig. 3. puted among the 16 character ratios for all HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 575

027 024

022 FEMALES o.s 036 031

@32

3E 37 330 2$ (33 0) 39

0-35 o 01Is 271 S 391 -85 . - *3 A 0. 4MALES 254

U 112 .5e

2.1 FACTOR ONEW 40,92% 22 A 6 9 4~Z

1 Fig. 7. Two-dimensional plots of factor means from the PCA for males

-.45 -.35 VECTOR ONE;45,2 5% - 15 -.05 for the males was influenced heavily by plastral shape (PW2/PL and PW3/PL) and the size of Fig. 6. Two-dimensional plots of the means of the canonical variates from the MANOVA-canonical two additional variables (V 1L/CL and GL/PL), analysis for females from all samples. whereas vector two was primarily influenced by plastron shape (PW1/PL, PW3/PL) and IASL/ PL. For the females, vector one was influenced by both shape (PW3/PL, FLL/PL, CW/CL) and samples and for both sexes. The first canonical size (V1L/CL and GL/PL) variables. Variation variate accounted for 37.27% and 45.25% of along vector two, for females, was influenced the variation among the samples for males and primarily by plastron shape (PW2/PL, PW3/PL) females, respectively; while the secopd canoni- and plastral scute size (IANSL/PL, IHSL/PL, cal variate expressed 23.88% and 21.93% of the IASL/PL). The major separation of the samples variation respectively. Plots of the means and in the MANOVA-canonical analyses appears to one standard deviation of the first two canoni- be on vector two, with samples of K.f arizonense cal variates for each sample are presented for and southern K. f flavescens tending to have a males (Fig. 5) and females (Fig. 6). narrower plastron and carapace, and a shorter Two major groups, separated by a wide hia- IANSL. tus can be discerned in Fig. 5 (males); samples of K. f. arizonense (34-38) and K. f. durangoense

(39) compose one group, and samples of K. f. 03024 flavescens (3-33) and K. f. spooneri (1-2) compose the other. Sample 1 (Illinois River K. f. spooneri) is separate from all samples of K.f. fa- 027034 027o20 *31 1 vescens. This gap, however, is bridged by Sam- ple 2 (Mississippi River K. f. spooneri), whose 0386 2e02 91 19 ellipse overlaps samples 4, 6, 26 and 29 (N = 1) 0 15 I s

of K. f. flavescens. Sample 10 (Neosho River) @a 1240*16 separates slightly along the second vector. S04040320 309 1e eo 3 Among the females (Fig. 6) the same two major groups are evident; samples of K. f. arizonense (34, 36-38) and K. f. durangoense (39-40) are separate from all other samples whereas sam- .~r PFEMALES ples of K. f. spooneri (1-2) show considerable overlap with those of K. f. flavescens (4, 6, 7, 9). The relative influence of each character ratio 4.2 0. FACTOR ONE; 38.55% 4 o0. on the first two canonical variates was deter- Fig. 8. Two-dimensional plots of factor means mined for both males and females. Vector one from the PCA for females. 576 COPEIA, 1982, NO. 3

1 2 6 26 4 MALES 3 FEMALES 8 8 9 .932 9 .906 23 12 12 13 13 15 16 14 14 25 17 16 25 17 15 18 7 32 32 33 22 10 26 19 18 29 28 20 31 22 33 28 20 31 27 24 5 27 11 11 24 38 19 34 21 35 37 38 39 37 34 36 36 40 1.987 1.737 1.487 1.237 0.987 0.737 0.487 0.237 39

Fig. 9. Distance phenogram 2.340 depicting 2.040 1.740 1.440 results1.140 0.840 0.540 of 0.240 the cluster analysis for males from all samples. Num- ber beneath the sex is the coefficient Fig. 10. Distance of cophenetic phenogram depicting the results correlation. of the cluster analysis for females from all samples. Number beneath the sex is the coefficient of cophenetic correlation. Factor one in the PCA separates samples of K. f. arizonense (34-38) and K. f. durangoense (39-40) from all other samples, for both males of K. f spooneri (1-2) cluster at a very low level (Fig. 7) and females (Fig. 8). Factor two tends with the two samples of K. f. favescens from to separate samples within the two major Nebraska (4, 6), and in the case of the males, groups, resulting in the slight separation of K.with sample 26 (Upper Rio Grande). f. arizonense samples (34-38) from samples of To analyze the relationship of K. f. spooneri K.f. durangoense (39-40). However, these anal- to K. f flavescens more effectively all multivari- yses fail to separate K. f. spooneri (1-2) from ateK. analyses were performed a second time, ex- f flavescens (3-33). Factors one and two for the cluding samples of K. f. arizonense and K. f du- males were influenced most heavily by (in de- rangoense. The relationships observed in the scending order) VIL/CL, IFSL/PL, PW3/PL, original analyses were not altered as a result of and FLL/PL, HLL/PL, IHSUPL, respectively; omitting these samples. Results from these sub- and IANSL/PL, GUPL, IFSUPL, and CW/CL, sequent analyses are therefore not figured or PW3/PL, PW2/PL, respectively, for the females. discussed. Ordination (such as PCA), and clustering may yield results which lead to different taxo- Color analysis.-The results of the color analysis nomic conclusions (Sneath and Sokal, 1973). A are presented in Fig. 11. All of the K.f. spooneri cluster analysis was therefore employed to ana- (locality 1) specimens are Class III (dark); how- lyze the data and serve as a comparison. Dis- ever, 70% of each of the two Nebraska localities tance phenograms depicting the phenetic re-(2, 3) are also Class III. No Class I (light) spec- lationships of all samples were computed for imens are found north of Kansas (locality 4), both males (Fig. 9) and females (Fig. 10) from which has 13.3% Class I specimens and 40.00% the distance matrix. Two major groups, iden- Class II (intermediate) specimens. Class III tical to those seen in the PCA, are evident from specimens occur less frequently south of Kan- this analysis. All samples of K.f. arizonense (33- sas. Localities 5, 6, 7 (Oklahoma, Texas and 38) and K. f durangoense (39-40) comprise one New Mexico, respectively) appear to be char- group, while all samples of K. f. flavescens and acterized by more light (40 to 60% Class I), and K.f spooneri make up the other group. Samples intermediate (20 to 50% Class II) specimens. HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 577

Although no Class I (light) specimens are found 105, 95* north of Kansas, at least one Class III (dark) SCALE specimen was found at all localities examined. 600 KM A distinct break in color variation is evident 42 10o between the Nebraska and Oklahoma localities, whereas the Kansas locality appears to be intermediate between them. Each of the three color classes are represented in localities 4, 5, 6, 7 (Kansas, Oklahoma, Texas and New Mex- ico, respectively) of Fig. 11. Dusky Brown to

Olive-Brown color gradation is evident from 30o 7B~ locality 1 of K. f spooneri to the New Mexico locality (7). ~Sso-

ZOOGEOGRAPHIC CONSIDERATIONS

Smith (1957) suggests that K. flavescens ex- panded its range eastward during a warm, post- Wisconsinan xerothermic period, in which a dry grassland habitat dominated much of the Fig. 11. Geographic distribution of the three col- prairie-peninsula region. With a subsequent or classes. White portion of pie represents percentage cooler, moister climate, a mesic forest invaded of Class I individuals; Stippled portion-Class II in- the region eliminating most of the grassland dividuals; Black portion-Class III individuals. Lo- habitat and the species associated with it. Some calities: 1. Mississippi River; N = 12; Class I = 0, Class II = 0, Class III = 12 (100%). 2. Nebraska of the more tolerant species (including K. fla- Sandhills; N = 17; Class I = 0, Class II = 5 (29.41%), vescens) were able to survive in relict sand prai- Class III = 12 (70.59%). 3. Republican River; N = ries of eastern Iowa, Illinois, and western In- 13; Class I = 0, Class II = 4 (30.77%), Class III = 9 diana. The disjunct populations of the mammal (69.23%). 4. Kansas; N = 15; Class I = 2 (13.33%), Geomys bursarius, the reptiles Tropidoclonion lin- Class II = 7 (46.67%), Class III = 6 (40.00%). 5. eatum, Heterodon nasicus, Pituophis melanoleucus, Oklahoma; N = 10; Class I = 6 (60.00%), Class II = Terrapene ornata and the amphibian Pseudacris 2 (20.00%), Class III = 2 (20.00%). 6. Texas; N = streckeri lend support to Smith's (1957) hypoth- 23; Class I = 14 (60.87%), Class II = 8 (34.78%), esis. According to this hypothesis K. f spooneri Class III = 1 (4.35%). 7. New Mexico; N = 12; Class has been isolated less than 4,000 years. I = 5 (41.67%), Class II = 6 (50.00%), Class III = 1 (8.33%). Sandhills K.f. favescens are known from early Pleistocene fossil material (Sand Draw Fauna), nearly 2.0 million years ago (Fichter, 1969; McGrew, 1944); just prior to the Pleistocene tundra-like vegetation (Flint, 1971). Braun glaciation. The Sandhills region was not gla- (1955) suggests that only taxa near the ice sheet ciated during the Pleistocene (Flint, 1959). Dur- were affected. Most of the Sandhills region was ing the maximum extent of Pleistocene glacia- sufficiently distant from all glacial advances to tion (Nebraskan and Kansas advances), glacial provide suitable habitat for a number of reptile ice only reached eastern Nebraska, whereas species, including K. flavescens. Barbour and Iowa, and all but extreme southern Illinois Schultz (1937) report two unidentified turtles were completely covered by glacial ice during in a fossil assemblage (Broadwater Fauna) of these advances (Flint, 1959). The ultimate de- Aftonian age (McGrew, 1944) from the south- velopment of the Sandhills region topography west edge of the Sandhills region (Morril Coun- occurred during the Pleistocene, when fluvial ty). The Broadwater Fauna is the most closely and pluvial processes eroded and reworked the related fauna to the Sand Draw Fauna (Mc- dune sand and the Ogallala beds. These factors Grew, 1944), and it is possible that the uniden- also contributed to the continued removal of tified turtles in the Broadwater Fauna are K. sand and loessal material from the Sandhills flavescens, a tolerant species. This would suggest region (Lugn, 1935, 1965). continuous habitation of the Sandhills region The direct cooling effects of the glacial by K.flavescens. ice The available climatic and fossil were probably limited to a narrow belt data along are consistent with this hypothesis. the edge of the ice sheet, characterized Although by a it cannot be determined with cer- 578 COPEIA, 1982, NO. 3 tainty when the Sandhills depends on one's K. interpretation flavescens of the subspe- became isolated, they are, at ciesthe concept. latest, Use of the as subspecies old trinomenas or older than the K. f spooneri has undergone populations, an interesting history, and and would a be expected to show wealth some of literature, degree both pro and ofcon, con-divergence. The fact that cerning no thestatistically use of the subspecies trinomen significant has variation is evident between either of these appeared (Systematic Zoology 1954-1956). populations and contiguous K. flavescens Somewould systematists feel justified in recognizing indicate isolation for a relatively short period any isolated group of populations as a distinct of time. This time period appears to be insuf-subspecies, and others recommend total rejec- ficient for significant evolutionary divergence tion of the subspecies category. In our opinion, to manifest itself to the point of recognizing a subspecies a should not be recognized unless it distinct taxon. Isolated populations of enhancesother the understanding of the biology of reptile species in the relict sand prairies the species,of not merely serving as a convenience Iowa, Illinois and Indiana (Tropidoclonion forlinea- cataloging museum specimens. Subspecies tum, Heterodon nasicus, Pituophis melanoleucus trinomens and should be applied when good evi- Terrapene ornata) have not been afforded dence of evolutionary divergence, as indicated unique subspecific status; nor has the isolated by phenetic breaks, exists. Mayr (1969, 1970) population of the mammal, Geomys bursarius defines a subspecies as "an aggregate of phe- (Smith, 1957; Conant, 1975). A relict amphib- notypically similar populations of a species in- ian, Pseudacris streckeri illinoensis, has been habiting rec- a geographic subdivision of the range ognized taxonomically (Smith, 1951, 1957) of but the species and differing taxonomically from Axtell and Haskell (1977) question its validity. other populations of the species." He goes on Kinosternon arizonense (=K. f. arizonense) to wasdescribe "differing taxonomically" as differ- described on the basis of fossil material of Plio- ing by diagnostic morphological characters. cene age (about 3.0 to 13.0 million years ago) Simpson (1961) suggests usage of the "75% from Arizona (Gilmore, 1922). This is also the rule"; "75% of the individuals of adjacent sub- earliest record of the genus Kinosternon, al-species will be unequivocally determinable," though the family dates back to the Oligocene thereby assuring significant differences in with Staurotypus (Mlynarski, 1976), and Xen- mean characters of subspecific populations. ochelys, an extinct kinosternid taxon (Ernst and The question concerning the taxonomic sta- Barbour, 1972). K. f. arizonense was probably tus of K. f. spooneri was approached by analyz- isolated in the Sonoran desert region at this ing the geographic variation of morphological time, or became isolated shortly thereafter features in K. flavescens in order to determine (possibly early Pleistocene). A possible northern whether a break occurs in any character or route (across the panhandle of New Mexico) characters between K. f spooneri and K. f. fa- for the invasion of the Sonoran desert region vescens. This is not inconsistent with the sub- by K. flavescens could be hypothesized on the specific concepts of Mayr and Simpson dis- basis of one specimen of K. f. flavescens from cussed above. west of the continental divide (Sample 29, Fig. K. f. arizonense shows significant morpholog- 2). The continental divide is also less than 1,400 ical divergence from K. f. flavescens in all anal- meters altitude between Animas and Hachitas, yses to support its subspecific recognition. It is New Mexico, well within the altitudinal range also separated from K. f durangoense by the of K.flavescens in New Mexico (Degenhardt and PCA. K. f. spooneri, however, shows very little Christiansen, 1974). K. f. durangoense is mor- morphological divergence. None of the multi- phologically very similar to K. f arizonense, de- variate or univariate analyses separated K. f. spite its geographic proximity to K.f. flavescens. spooneri from K. f flavescens. A case for the rec- Iverson (1979b) suggests that this morpholog- ognition of K. f. spooneri could not be made ical similarity may be due to convergent adap- from the color analysis. This character shows tation to similar arid environments, rather than clinal variation and only 30 to 40% of the spec- common descent. imens of K. f. spooneri (samples 1-2) can be dis- tinguished from northern K. f. favescens (sam- TAXONOMIC CONCLUSIONS ples 4, 6) using this character. Both Smith (1951) and Iverson (1979b) list key characters Ultimately, the decision concerning the taxo- in distinguishing K. f spooneri from K. f. flaves- nomic status of the subspecies of K. flavescens cens (among them, a longer gular scute, a short- HOUSEAL ET AL.-KINOSTERNON FLAVESCENS GEOGRAPHIC VARIATION 579

er inter-humeral seam and darker coloration in cisms, or discussion of the data: J. R. Dixon, D. K. f. spooneri), which Iverson (1979b) used J. in Schmidly, I. F. Greenbaum, M. D. Engstrom, the key to subspecies of K.flavescens. This study J. W. Sites, Jr., F. L. Rose, J. B. Iverson, C. J. reveals considerable overlap of these charac- McCoy, R. C. Vogt, J. L. Christiansen, E. O. ters, and others, between K. f. flavescens and Moll,K. G. D. Schnell, H. M. Smith and J. M. In- f. spooneri. glis. Chet Churchill and Francis Rose aided in When all of the analyses (multivariate, uni- the field work. Robert Ellis granted permission variate and color) are considered together, a to collect on the Valentine National Wildlife Ref- clear and relatively consistent pattern of geo- uge. Daniel Gallagher assisted in examination of graphic variation emerges. K. f. arizonense is some specimens. John Iverson was generous in distinct from K. f. flavescens, although clearly providing unpublished manuscripts, access to these forms are closely related and conspecific. data, and advice on collecting localities. K. f. arizonense and K. f. durangoense are mor- We thank the following for providing valu- phometrically very similar but show slight di- able information and/or making material avail- vergence in the PCA. K.f. spooneri, however, is able for examination: John Applegarth, Walter inseparable from the Nebraskan samples (4, 6) Auffenberg, Charles Carpenter, James Chris- of K.f.flavescens. Perhaps K.f. spooneri is a valid tiansen, William Degenhardt, William Duell- subspecies but includes populations from Ne- man, John Iverson, Donald Kangas, Arnold braska. If this is true then a zone of intergra- Kluge, Charles Lowe, John Lynch, Hymen dation occurs in Kansas, and this is somewhat Marx, C. J. McCoy, Leonard McDaniels, John supported by the color analysis (Fig. 11). Of the Mecham, Edward Moll, Paul Neiswonger, Ron- morphometric analyses, the cluster analysis sup- ald Nussbaum, Francis Rose, Gary Schnell, ports inclusion of the Nebraskan samples with Robert Semmler, Daniel Shorter, Donald Tin- K. f. spooneri (Figs. 9, 10) but the level at which kle, Dick Vogt, Ernest Williams, George Zug these samples cluster is so low that several other and Richard Zweifel. clusters would also have to be recognized tax- This manuscript is part of a thesis submitted onomically. No identifiable divergence between by TWH in partial fulfillment of the require- K. f flavescens and K. f. spooneri is evident in the ments for the MS degree from Texas A&M results of the PCA. In the MANOVA-canonical University. analyses (Figs. 5, 6) and univariate analyses

(Fig. 3, 4), the samples of K. f. spooneri overlap LITERATURE CITED those of Nebraskan K. f. flavescens but there is no separation of these samples (1-6) from AGASSIZ, the L. 1857. Contributions to the Natural His- other samples of K. f. flavescens. Although tory we of the United States of America. Vols. 1, 2. recognize this pattern of geographic variation, Little, Brown and Co., Boston. no sample or combination of samples are ALBRECHT, suf- G. H. 1978. Some comments on the use ficiently distinct to warrant recognition as a oftax- ratios. Syst. Zool. 27:67-71. on distinct from K. f.lavescens. We therefore ATCHLEY, W. R. 1978. Ratios, regression, intercepts, recommend that K. f. spooneri be placed in andsyn- the scaling of biological data. Ibid. 27:78-83. --, AND D. ANDERSON. 1978. Ratios and the sta- onymy with K. f. flavescens. tistical analysis of biological data. Ibid. 27:71-78. --, C. T. GASKINS AND D. ANDERSON. 1975. Sta- ACKNOWLEDGMENTS tistical properties of ratios in biological data. Amer. Zool. 15:829. We are grateful to LGL Ecological Research , AND . 1976. Statistical proper- Associates, Inc., and all of their employees who ties of ratios. I. Empirical results. Syst. Zool. worked on this project during the summer of 25:137-148. 1979. We thank Benny J. Gallaway of LGL, for AXTELL, R. W., AND N. HASKELL. 1977. An interhiatal cooperation and assistance throughout the population of Pseudacris streckeri from Illinois with course of this study and Will Carpenter of an assessment of its postglacial dispersion history. Natur. Hist. Misc., Chicago Acad. Sci. 202. Monsanto for providing the opportunity to do BARBOUR, E. H., AND C. B. SCHULTZ. 1937. An early this research. Funding for this project was pro- Pleistocene fauna from Nebraska. Amer. Mus. vided by Monsanto Agricultural Products Co., Novitates 942. contracted through LGL Ecological Research BERRY, J. F. 1978. Variation and systematics in the Associates, Inc. Kinosternon scorpiodes and K. leucostomum complexes The following people offered advice, criti- (Reptilia: Testudines: Kinosternidae) of Mexico 580 COPEIA, 1982, NO. 3

and Central America. mud Unpubl. turtle, Kinosternonflavescens PhD diss.,(Testudines: Kin-Univ. Utah. osternidae). Ibid. 212-225. BRAUN, E. L. 1955. Phytogeography of unglaciated - , AND J. F. BERRY. 1979. The mud turtle genus eastern United States and its interpretation. Botan. Kinosternon in northeastern Mexico. Herpetologica Rev. 21:297-375. 35:318-324. BROWN, L. E., AND D. MOLL. 1978. The status of LUGN, the A. L. 1935. Pleistocene geology of Nebraska. nearly extinct Illinois mud turtle with recommen- Nebraska Geological Survey Div., 10. dations for its conservation. Milwaukee Pub. Mus. . 1965. The origin of loesses and their relation Spec. Publ. Biol. Geol. No. 3. to the Great Plains in North America, p. 139-182. CAHN, A. R. 1931. Kinosternonflavescens: a surprising In: Loess and related eolian deposits of the world, turtle record from Illinois. Copeia 1931:120-123. vol. 12. C. B. Schultz and J. C. Frye (eds.). Proc. --. 1937. The turtles of Illinois. Ill. Biol. Monog. VIIth Congr., Internat. Assoc. Quaternary Res. 35. Univ. Nebraska Press. CONANT, R. 1975. A field guide to the reptiles MAYR, and E. 1969. Principles of systematic zoology. amphibians of eastern and central North America. McGraw Hill Book Co., New York. Houghton Mifflin Co., Boston. . 1970. Populations, species and evolution. , AND J. F. BERRY. 1978. Turtles of the family Belknap Press of Harvard Univ. Press, Cambridge. Kinosternidae in the southwestern United States MCGREW, P. O. 1944. An early Pleistocene (Blancan) and adjacent Mexico: identification and distribu- fauna from Nebraska. Field Mus. Nat. Hist. (Chi- tion. Amer. Mus. Novitates 2642. cago), Pub. 546, Geol. Ser. 9:33-36. CORRUCCINI, R. S. 1977. Correlation properties MLYNARSKI,of M. 1976. Testudines, p. 1-130. In: ratios. Syst. Zool. 26:211-214. Handbuch der Paleoherpetologie. Oskar Kuhn DEGENHARDT, W. G., AND J. L. CHRISTIANSEN. 1974. (ed.). Gustav Fischer Verlag. Distribution and habitats of turtles in New Mexico. ROHLF, F. J. 1968. Stereograms in numerical taxon- Southwest Nat. 19:21-46. omy. Syst. Zool. 17:246-255. DODSON, P. 1978. On the use of ratios in growth - , AND J. KISHPAUGH. 1972. Numerical taxon- studies. Syst. Zool. 27:62-67. omy system of multivariate statistical programs. DUELLMAN, W. E., T. FRITTS AND A. E. LEVITON. SUNY, Stony Brook, New York. 1978. Museum acronyms. Herp. Rev. 9:5-9. SERVICE, J. 1972. A user's guide to the statistical anal- ERNST, C. H., AND R. W. BARBOUR. 1972. Turtles of ysis system. North Carolina State Univ., Raleigh. the United States. Univ. Press Kentucky, Lexing- SIMPSON, G. G. 1961. Principles of animal taxonomy. ton. Columbia Univ. Press, New York. FICGHTER, L. S. 1969. Geographical distribution SMITH, andP. W. 1951. A new frog and a new turtle from osteological variation in fossil and recent specimens the western Illinois sand prairies. Bull. Chicago of two species of Kinosternon (Testudines). J.Acad. Her- Sci. 9:189-199. petol. 3:113-119. - . 1957. An analysis of post-Wisconsin biogeog- FLINT, R. F. 1959. Glacial map of the United raphy States of the prairie peninsula region based on dis- east of the Rocky Mountains. Scale 1:1,750,000. tributional phenomena among terrestrial verte- Geol. Soc. Amer., New York. brate populations. Ecology 38:205-218. . 1971. Glacial and quaternary geology. John SMITHE, F. B. 1975. Naturalist's color guide. Amer. Wiley and Sons, New York. Mus. Nat. Hist., New York. GILMORE, C. W. 1922. A new fossil turtle, Kinsternon SNEATH, P. H. A., AND R. R. SOKAL. 1973. Numerical arizonense, from Arizona. Proc. U.S. Nat. Mus. taxonomy. W. H. Freeman and Co., San Francisco. 62:1-18. HARTWEG, N. 1938. Kinosternon favescens stejnegeri, DEPARTMENT a OF WILDLIFE AND FISHERIES SCI- new turtle from northern Mexico. Occ. Pap. ENCES,Mus. TEXAS A&M UNIVERSITY, COLLEGE Zool. Univ. Mich. 371. STATION, TEXAS 77843 AND LGL ECOLOGICAL HILLS, M. 1978. On ratios-A response to Atchley, RESEARCH ASSOCIATES, INC., 1410 CAVITT, Gaskins, and Anderson. Syst. Zool. 27:61-62. BRYAN, TEXAS 77801. PRESENT ADDRESS IVERSON, J. B. 1978. Distributional problems of the genus Kinosternon in the American southwest. (TWH): Co- DEPARTMENT OF BIOLOGY, TEXAS peia 1978:476-479. A&M UNIVERSITY, COLLEGE STATION, TEXAS --. 1979a. On the validity of Kinosternon arizo- 77843; (MDS): 12508 ROSEMONT NE, ALBU- nense Gilmore. Ibid. 1979:175-177. QUERQUE, NEW MEXICO 87112. Accepted 28 -~ . 1979b. A taxonomic reappraisal of the yellow May 1981.