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Scientific Papers QL 638 .P4 S53 1999 Scientific Papers Natural History Museum The University of Kansas Number 12:1-16 28 July 1999 A Reexamination of the Phylogenetic Relationships of the Sand Darters (Teleostei: Percidae) By Kate A. Shaw', Andrew M. Simons^ and E. O. Wiley' 'Division oflchthyologi/, Natural Histonj Museum, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-2454, USA ami ^Bell Museum of Natural History and Depmrtment of Fisheries and Wildlife, The University of Minnesota, 1980 Fohvell Avenue, St. Paul, Minnesota 55108-6124, USA CONTENTS ABSTRACT 1 INTRODUCTION 2 Acknowledgments 3 MATERIALS AND METHODS 3 RESULTS 6 DISCUSSION 10 LITERATURE CITED 12 APPENDIX 1. Specimens Examined 13 APPENDIX 2. Biochemical Data 14 APPENDIX 3. Character Matrix 15 ABSTRACT Phylogenetic relationships among the Boleosoma group of darters were examined for allozymic variation at 25 presumptive gene loci alone and in combination with 29 morphological char- acters. Qualitative analyses of allozymic and morphological— variation resulted in two most parsimoni- ous trees showing the following relationships: Tree 1. (Etheostoma davisoni ((£. longimanum, E. nigrum)— (E. vitreum (£. clarum (£. pellucidum (E. meridianuni (E. vivax (E. bifascia, E. beanii)))))))); Tree 2. (E. davisoni (E. nigrum (E. longimanum (E. vitreum (E. clarum (E. pellucidum (E. meridianuni (E. z'iz'ax (E. bifascia, E. beanii))))))))). Quantitative analyses that incorporate an estimate of allelic frequencies were © Natural History Museum, The University of Kansas ISSN No. 1094-0782 QL 638 .P4 S53 1999 Scientific Papers Natural History Museum The University of Kansas Number 12:1-16 28 July 1999 A Reexamination of the Phylogenetic Relationships of the Sand Darters (Teleostei: Percidae) By Kate A. Shaw', Andrew M. Simons^ and E. O. Wiley' ^Division Natural and of Ichthyology , History Musciiin, Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-2454, USA and 'Be// Museum of Natural History and Department of Fisheries and Wildlife, The University of Minnesota, 1980 Fohvell Avenue, St. Paul, Minnesota 55108-6124, USA CONTENTS ABSTRACT 1 INTRODUCTION 2 Acknowledgments 3 MATERIALS AND METHODS 3 RESULTS 6 DISCUSSION 10 LITERATURE CITED 12 APPENDIX 1. SPEaMENS Examined 13 APPENDIX 2. Biochemical Data 14 APPENDIX 3. Character Matrix 15 ABSTRACT Phylogenetic relationships among the Boleosoma group of darters were examined for allozymic variation at 25 presumptive gene loci alone and in combination with 29 morphological char- acters. Qualitative analyses of allozymic and morphological variation resulted in two most parsimoni- . — ous trees showing the following relationships: Tree 1 (Etheostoma davisoni ((£. lon^imanum, E. nigrum) (E. vitreum (£. claruin (£. peltucidum (£. meridiainim (£. znz'ax (£. hifascia, E. beanii)))))))); Tree 2. —(£. davisoni (E. nigrum (£. longimanum (£. vitreum (£. clarum {£. pellucidum (£. meridianum {£. vivax (E. hifascia, E. beanii))))))))). Quantitative analyses that incorporate an estimate of allelic frequencies were © Natural History Museum, The University of Kansas ISSN No. 1094-0782 ScFENTiFic Papers, Natural History Museum, The University of Kansas used to discriminate among trees and showed that Tree 1 was 0.598 FREQPARS units shorter than Tree 2. There was both congruence and complementarity in the support provided by allozymic and mor- phological datasets. The sister-group relationship and allopatric distributions of £. beanii and £. bifascia are consistent with their origin resulting from vicariant speciation associated with the origin of the Mobile Basin. Key words: Etheostoma, Ammociypta, allozymes, historical biogeography, phylogeny INTRODUCTION The sand darters are slender, translucent elongate, oi Ammocrypta within the genus Etheostoma and suggested darters known for their habit of themselves in burying that Etheostoma (loa) vitreum was the sister group to the substrates. Six of sand darters are sandy species currently subgenus Ammocrypta. This change resulted in four sub- classified in the Etheostoma subgenus Ammocri/pta: genera being included in the Boleosoma group of Etheostoma: E. beanii, E. vivax, E. claruni, E. and £. peUucidum, bifascia, Boleosoma, loa, Vaillantia, and Ammocrypta. Simons' hypoth- meridianum. esis of relationships within the subgenus Ammocrypta dif- The taxonomic history of Ammocrypta began with its fered somewhat from those postulated by Williams (1975). use as a generic name by Jordan (1877) with his descrip- Simons (1992) did find support for the £. peUucidum group; tion of y4. beanii. Three additional species now included in £. meridianum and £. peUucidum were sister taxa, and £. the subgenus Ammocrypta (originally described as vivax was the sister group to this pair. However, Simons Pleurolepis pellucida, A. vivax, and A. clam) were described (1992) did not find support for the £. beanii group; instead, by 1886 (Agassiz, in Putnam, 1863; Hay, 1883; and Jordan the species pair £. beanii-E. bifascia was the sister group to and Meek, 1885, respectively). Bailey and Gosline (1955) the £. peUucidum group and £. clarum was the sister group added Crystallaria asprelln to the genus Ammocrypta, but to all other members of the subgenus Ammocrypta. it in a placed monotypic subgenus Crystallaria. Although The relationships hypothesized by Simons (1989, 1991, and Gosline (1955) for a classification of Bailey argued 1992) and based on phylogenetic analyses, have not been darters three (Percina, and containing genera Anunocn/pta, widely accepted. For example, Etnier and Starnes (1993), Etheostoma) that was a few authors generally accepted, (e.g., Jenkins and Burkhead (1994), Mettee et al. (1996), and Moore, 1968; Miller and Robison, 1973) continued to rec- Pflieger (1997) all continued to recognize the genus as a Williams (1975) ognize Crystallaria monotypic genus. Ammociypta in their state guides (Tennessee, Virginia, Ala- described two additional within the species subgenus bama, and Missouri, respectively). Although Jenkins and Within this his A. beanii con- Ammocrypta. subgenus, group Burkliead (1993) alone justified their retention of the ge- sisted of the with a few scale rows (£. beanii, E. species nus Ammocrypta, in no case is an explicit alternative phy- and £. clariim) and his A. contained bifascia, pellucida group logenetic analysis or hypothesis presented. Wood and the to almost scaled (£. partially completely species Mayden (1997) however, found support for a sister-group E. meridianum, and £. vivax). peUucidum, relationship between Etheostoma beanii and Crystallaria in Williams (1975) considered the genus Ammocrypta maximum parsimony analyses and in their most parsi- more closely allied to the subgenus hnostoma of the genus monious FREQPARS tree, but they did not make any taxo- Percina than to Etiieostoma. However, Page and Whitt nomic recommendations based on this result. In the absence (1973a) argued that Percina was monophyletic and sug- of an explicit alternative phylogenetic analysis of more than gested that Ammocrypta was related to Etheostoma; this con- two taxa, we conrinue to consider members of Ammocrypta, clusion was based on a unique LDHB4 isozyme found only members of the genus Etheostoma, subgenus Ammocn/pta. in Perciim. (Their assessment of the of this charac- polarity The particular relationships of these species are of in- ter is unclear— in their 4, Etheostoma and Fig. p. 6, terest to biogeographers. Wiley and Mayden (1985) fol- are united a derived LDH B4 Ammocrypta by isozyme.) Page lowed the relationships suggested by Williams (1975) and and Whitt (1973b) stated that the TO shared £. isozyme by postulated that the distribution of members of the chlorosoma and A. indicated that (Vaillantia) pellucida Etheostoma beanii group was the result of a western was more related to Etheostoma than Ammocrypta closely vicariance event loosely associated with the Mississippi to Percina. River. Etheostoma clarum is found in the Mississippi River Simons (1989, 1991) removed Crystallaria from Basin and drainages to the west, and the presumed sister Ammocrypita and hypothesized that the phylogenetic posi- taxon (£. beanii-E. bifascia) is found east of the Mississippi tion of Crystallaria was basal to Etheostoma and Percina. River mainstem. Wiley and Mayden (1985) also suggested Simons (1989, 1992) subsumed the remaining six species that the distributions of two other groups could be attrib- Phylogenetic Relationships of Sand Darters uted to vicariance events involving the Mobile Bay Basin. Acknowledgments Ethcostoina beaiiii is found in the Mobile Basin and west- We thank G. Harp (Arkansas State University), L. Page, ward to the Mississippi River mainstem, whereas E. bifascia P. Ceas, and C. Johnston (Illinois Natural History Survey), is only found east of Mobile Bay. A similar situation is F. Pezold (Northeast Louisiana University), R. Jenkins found in the £. pelliicidiim group. Etheostoma meridiaiiiim is (Roanoke College), H. Bart and M. Taylor (Tulane Univer- a Mobile Basin whereas E. and E. vivax endemic, pellucidum R. sity), R. Mayden (University of Alabama), Cashner and are found in north and west of the Mobile Ba- drainages F. T. J. Grady (University of New Orleans), and Cross, sin. Simons' (1992) results did not substantiate the Schmidt, D. Siegel-Causey, and K. Toal, III (University of vicariance event associated with the Mississippi River or Kansas) for their help collecting specimens used in this a vicariance event
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