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The American Percid Fishes of the Subgenus Villora Bruce B

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The American Percid Fishes of the Subgenus Villora Bruce B ,n 3k THE AMERICAN PERCID FISHES OF THE SUBGENUS VILLORA BRUCE B. COLLETTE, Ichthyological Laboratory, Fish and Wildlife Service U.S. National Museum, Washington 25, D.C. and RALPH W. YERGER, Department of Biological Sciences, Florida State University, Tallahassee, Florida Reprinted from TULANE STUDIES IN ZOOLOGY Vol. 9, No. 4, pp. 213-230 March 30, 1962 THE AMERICAN PERCID FISHES OF THE SUBGENUS VILLORA BRUCE B. COLLETTE, Ichthyological Laboratory, Fish and Wildlife Service U.S. National Museum, Washington 25, D.C. and RALPH W. YERGER, Department of Biological Sciences, Florida State University, Tallahassee, Florida I. INTRODUCTION with those for the species of Hololepis in The aims of this paper are threefold: to another paper (Collette, 1962, Tables 38- define the subgenus Villora, to resurrect and 49 ). define Etheostoma (Villora) okaloosae ( Fowler ), and to examine intra-specific II. ACKNOWLEDMENTS variation in E. (Villora) edwini. Villora The section on Etheostoma edwini is was described as a new genus by Hubbs based on part of a doctoral dissertation pre- and Cannon ( 1935 ) on the basis of the sented to Cornell University by the senior new species edwini. Bailey (1951), in Bailey author. Professor Edward C. Raney, who Winn and Smith ( 1954), and in Bailey and served as chairman of the committee, pro- Gosline ( 1955) reduced the many nominal vided, in addition to constant interest, sup- genera of darters to three: Percina, Am- port for the study through National Sci- mocrypta and Etheostoma. In 1955 (Fig. 1), ence Foundation Grants 2893 and 9038. Bailey and Gosline arranged the darters in The section on E. okaloosae is part of a order of increasing specialization and some study on the fishes of the Florida panhandle former genera were utilized as subgenera. undertaken by the junior author with sup- Etheostoma edwini was listed under the port from a National Science Foundation subgenus Hololepis„ without further com- Grant G-6260 and from the Research ment. We cannot subscribe to this view. Dur- Council of the Florida State University. The ing his study of the subgenus Hololepis, the many contributions to this project by Ardith senior author (Collette, 1960) became B. Cochran are gratefully acknowledged, as convinced that Villora should be regarded as are the services rendered by W. Bruce Wal- distinct from Hololepis. Through extensive den and Robert F. Christensen, graduate stu- collecting in western Florida, the junior dents at Florida State University. Colonel author (Yerger, 1960) discovered that John N. Reynolds, Commander, Eglin Air Etheostoma okaloosae ( Fowler) is a valid Force Base, Florida, granted permission to species referable to the subgenus Villora, collect on the military reservation and ex- and concluded that it must be removed from tended many courtesies. James E. &hike the synonymy of E. swami ( Jordan) where compared the type of E. okaloosae with sev- it was placed by Bailey, Winn and Smith eral specimens sent to him by the junior (1955). author. The methods of this study are similar to We wish to express our appreciation to those of Collette (1962). However, here the following persons and the institutions we present the counts of the unpored they represent for having loaned specimens ( Table 3) as well as the pored and total under their care or for making available lateral-line scales. This character is unim- facilities at their institutions: Reeve M. portant in the subgenus Hololepis, but Bailey, University of Michigan Museum of proves roost useful in Villora. Measure- Zoology (UMMZ) ; James E. Bohlke, Acad- ments were made following the technique emy of Natural Sciences of Philadelphia described by Hubbs and Cannon (1935). In ( ANSP) ; Robert H. Gibbs, Jr., Boston the course of this study, more than 1100 University ( BU); A. Frederick Hemphill, specimens of E. edwini and nearly 200 of Spring Hill College, Alabama (UAIC) ; E. okaloosae were examined. For purposes Robert H. Kanazawa, Ernest A. Lachner, of comparison, summary counts for the two and Leonard P. Schultz, United States Na- species of Villora are being presented along tional Museum (USNM) ; E. E. Prather, 214 Tulane Studies in Zoology Vol. 9 Alabama Polytechnic Institute (API); Ed- base; dark humeral spot behind the opercle ward C. Raney, Cornell University (CU); and above the pectoral fin origin usually Royal D. Suttkus, Tulane University (TU); prominent; breeding tubercles absent; males use was also made of specimens in the reaching a larger maximum size than fe- Florida State Museum, University of Flori- males; habitat moderately fast, clear streams da (UF), and in Florida State University with patches of vegetation; range northern (FSU). and western Florida and below the Fall Line Many people have assisted by discussing in southwestern Georgia and in Alabama. our study with us or by reading the manu- Hubbs and Cannon (1935) placed Villora script and offering valuable suggestions. In between Hololepis and the catch-all Poeci- particular we wish to thank Reeve M. lichthys. They felt that many of the features Bailey, Daniel M. Cohen, Carl L. Hubbs of Villora were primitive and suggested and Edward C. Raney. Douglass M. Payne, that it may have evolved from an ancestor Cornell University, took the photographs of Hololepis. The relationships of Villora, (Figs. 1-6). Miriam W. Schaffer drew the however, do not seem to lie with Hololepis figures of the genital papillae. Mildred H. (see Collette, 1962). The rediscovery of Carrington completed one of the maps (Fig. E. okaloosae, which is obviously closely re- 10). Sara E. Collette and Alice B. Holland lated to E. edwini and more different from typed several drafts of the manuscript. Hololepis than edwini, gives support to our view that Villora should stand as a sub- genus separate from Hololepis. In a num- III. SUBGENUS Villora ber of characters E. okaloosae is somewhat Villora Hubbs and Cannon, 1935:11-13 intermediate between E. edwini and E. (Oli- (type species Villora edwini Hubbs and gocephalus) swami ( Jordan). There seems Cannon, 1935, by original designation). to be a trend toward a shorter genital papilla Lateral line slightly arched upward, in- (Fig. 7), a more highly arched lateral line, complete to complete; pored lateral-line a slenderer caudal peduncle, and fewer pored scales 20 to 36, unpored 0 to 18, total 32 lateral-line scales as one moves from E. (O.) to 42; infraorbital canal always complete, swami through E. (V.) okaloosae to E. (V.) usually with 8 pores; supratemporal canal edwini. In the last three characters the evo- usually complete in adults; coronal pore lutionary trend parallels that of the sub- present; interorbital pores usually 2; pre- genus Hololepis, resulting in a superficial operculomandibular pores usually 9 or 10; similarity between the Villora and Hololepis vomer and palatine toothed; preopercle en- lines. Villora differs from the subgenus tire or crenulate; branchiostegal membranes Hololepis by lacking breeding tubercles and separate or narrowly conjoined; branchios- by having a nearly complete lateral line, ex- tegal rays usually 6; breast naked to entirely tending past the origin of the second dorsal covered with scales; nape, preopercle, and fin (occasionally complete in E. okaloosae); opercle completely covered with imbedded a much less arched lateral line; the genital cycloid to exposed ctenoid scales; parietal papilla in the form of a low tube crowned and interorbital regions naked; belly cov- with villi (Collette, 1962: Fig. 1); and a ered with unmodified ctenoid scales; flesh dark humeral spot. The species live in clear, opaque; body rather stocky and somewhat moderately fast-flowing, sandy-bottomed compressed; vertebrae 33 to 38; premaxil- streams. Villora differs from many of the lary frenum broad; first dorsal fin mod- species of the subgenus Oligocephalus in erately high, with 8 to 11 spines that lack having a somewhat arched lateral line and thickened fleshy tips; second dorsal fin in the shape of the genital papilla. We somewhat higher than first dorsal, with 9 have not studied thoroughly the differences to 13 rays; anal spines usually 2, the first between Oligocephalus and Villora. This thicker than the second; anal soft-rays 5 to matter deserves further study. 9; pectoral rays usually 12 or 13; segmented caudal rays: upper 8 to 13, lower 7 to 10, IV. SPECIES OFVillora total 15 to 22; genital papilla of breeding The two species that we refer to the sub- female a low tube crowned with villi; pelvic genus Villora differ in a number of respects fins closely approximated, separated by a ( Table 1) . Frequency distributions of the distance equal to about one-half the pelvic more important characters are presented in No. 4 Collette and Yerger: Subgenus Villora 215 Tables 2-8. Table 9 compares the propor- lack distinct basi-caudal spots; some have tional measurements of the two species. blackish median spots and vestiges of dorsal and ventral spots. Eight to twelve lateral Etheostoma (Villora) edwini blotches are sometimes present along the ( Hubbs and Cannon) sides; they may extend down to the under- Villora edwini—Hubbs and Cannon, side of the caudal peduncle. There are 1935:13-16 (original description) ; Carr, usually about nine dark brown dorsal sad- 1937:83 (Fla.); Fowler, 1941:242, fig. 13 dles. Melanophores on the sides below the ( not 3 as given) (Fla.) ; Bailey and Hubbs, lateral line are generally arranged in a 1949:10 (edwini one of a group of en- prominent X-pattern as described by Hubbs demic species, probably relicts, in sinkhole and Cannon ( 1935). Chromatophores be- region of N. Fla.). low the lateral line are mostly black; those above the lateral line are brown. The gen- Etheostoma edwini—Bailey, Winn, and ital papilla is usually devoid of pigment, Smith, 1954:144, 156 (range extension W but the dorsal surface of the distal portion to Perdido R., and N into S Ala.); Bailey is sometimes pigmented.
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