Fishes of Texas Project Documentation

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Since scientific names of species change over time and data sources vary in adherence to standard names, we synonymized all invalid names to a standard taxonomy. However, researchers have noted the presence of fish populations that appear to be unique and often describe them as a subspecies or a unique form of some sort, but lack formal description or even a name. Usually they are smaller populations and geographically isolated from a larger sister taxa population, but because of that isolation they are on their own evolutionary trajectory and thus different or can become different from the larger population and worth knowing about. To maximize awareness and potential conservation, we aim to recognize these various forms, even before they are formally described as species - some, of course, may never be formally described. Thus we prefer to think of our lowest level taxa names as Operational Taxonomic Units.

FoTX Standard Taxonomy

Our taxonomy and vernacular (common) names conform to the American Fisheries Society' common names for fishes publication (Page et al., 2013) when possible. Some come from taxa noted in the Hubbs et al. checklist and key (2008), the American Fisheries Society's 2008 report on Conservation Status of Imperiled North American Freshwater and Diadromous Fishes (Jelks et al., 2008) and others are Operational Taxonomic Units we justify below. Some taxa names were received from our donors for species not known to occur in North America, and thus are not included in these publications that focus only on North America, but are none-the-less valid taxa names. For these, our standard is the Global Biodiversity Information Facility's (GBIF) Backbone Taxonomy which draws from the Catalog of Life, the Integrated Taxonomic System, and others.

Synonymization to FoTX Standard

Focusing primarily on georeferenced records occurring in our study area (recall we also have records that fall outside our study area), each taxon in the database was assigned a synonym from our accepted taxonomy. Sometimes names were not provided by the data provider, and in other cases synonymy was not possible because the taxon was unrecognized or not identified to the species level. In other records generic names and species names were each recognizable, but when paired (e.g., Lepisosteus humilis) did not constitute a valid binomial taxonomic name. The California Academy of Science's Catalog of Fishes proved invaluable for researching names. Fishbase, and the Integrated Taxonomic System (ITIS) were useful resources as well. For data sets processed after tracks 1 and 2 we relied heavily on a tool provided by GBIF for synonymizing names to the GBIF Backbone Taxonomy but, also had to manually research some names.

All records that could not be unambiguously synonymized to our standard taxonomy for any reason, including records for specimens that lacked determinations entirely were flagged for specimen examination.

Taxa-Specific Decisions

Some taxa required specific treatments following recommendations in published literature and other sources. Some are controversial in that very little evidence exists supporting their being distinct forms and some may turn out not to be distinct forms. We err on the side of recognizing these operational taxonomic units in hopes that recognition will result in further research to resolve their status. The following taxa listed in bold, and arranged alphabetically by genus and species, are recognized by the Fishes of Texas Project as explained. Many of these are also adopted by the American Fisheries Society' common names for fishes publication (Page et al. 2013), but not all. Note that when specimen determinations are based on published biogeography, they have been applied to our georeferenced records only.

Ambloplites rupestris: A disjunct and likely introduced population of Ambloplites persists in central Texas. Specimens of this population have sometimes been called A. ariommus (Page and Burr 2011) and our examination of some suggests that they may be hybrids. More work is needed, but presently we follow Brown (1953), who determined, based on stocking records, that these were most likely A. rupestris. Effective in version 1.0.

Astyanax argentatus: Schmitter-Soto (2017) places the Texas and northern Mexico Astyanax, previously A. mexicanus, as A. argentatus. Effective in version 3.0.

Baldwinella aureorubens: Hemanthias aureorubens is now Baldwinella aureorubens according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Baldwinella vivanus: Hemanthias vivanus is now Baldwinella vivanus according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Carpiodes sp (Llano River Carpsucker): Some specimens of Carpiodes from the upper and middle Colorado River appear to be distinctively elongated compared to more typical-looking Carpiodes carpio found in the Colorado River. Suttkus and Bart (2002) and Chen et al. (2009) pointed out this distinctness and also recognized a similar distinct form occurring in the Rio Grande drainage. Bart (pers. comm. 2015) continues work describing and defining distributions of Texas forms of Carpiodes. We await the outcome of continued research, but will not be surprised if the slender form from the Colorado eventually is found to represent a distinct species. Many of these elongated specimens have been previously called Carpiodes cyprinus (Version 1), since they key out closely to them. They differ, primarily in body and head shape, from both C. carpio and C. cyprinus. We recognize this distinct form and expect many of the specimens now called C. carpio in the Colorado River to be re-determined to this form. Effective in version 3.0.

Chasmodes longimaxilla: Williams (1983) and McEachran and Fechhelm (2010) agree that Texas specimens referred to Chasmodes bosquianus are instead C. longimaxilla. Effective in version 2.0.

Choranthias tenuis: Anthias tenuis is now Choranthias tenuis according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Coptodon zillii Tilapia zilli is now Coptodon zillii based on genetic analysis by Dunz and Schliewen (2013). This change had been adopted in Fishbase and iNaturalist as well. Effective in version 3.0.

Cycleptus sp (Rio Grande Blue Sucker): Although undescribed, we follow Burr and Mayden (1999), Buth et al. (2001) and Bessert (2006) in referring all records of this genus from the Rio Grande drainage to this undescribed species. Effective in version 2.0.

Cyprinella: Texas Cyprinella are mostly morphologically similar and, when range overlap is not indicated, determinations here were mostly done on the basis of ranges defined in Schönhuth and Mayden (2010). Effective in version 3.0.

  • Cyprinella camura: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella lutrensis we must examine specimens and can't make determinations based solely on geography.  Effective in version 3.0.
  • Cyprinella panarcys: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella sp 1 (Rio Conchos Shiner) we must examine specimens and can't make determinations based solely on geography.  Effective in version 3.0.
  • Cyprinella proserpina: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella suavis and Cyprinella sp 1 (Rio Conchos Shiner) we must examine specimens and can't make determinations based solely on geography.  Effective in version 3.0.
  • Cyprinella lepida: We follow Richardson and Gold (1995) and Schönhuth and Mayden (2010) and refer all Cyprinella lutrensis from the Frio River upstream of Uvalde to C. lepida. Effective in version 2.0.
  • Cyprinella lutrensis blairi: We resurrect this subspecies first described by Hubbs (1940). We identified records of this form by inference based on published literature, which places the subspecies in Brewster County at Garden Spring (also known as Monument Spring) and in Pena Colorad(a/o) Creek (1940 Hubbs' description) and in Maravillas Creek (Miller et al. 1989). We apply this subspecies name to all records identified as C. lutrensis from these locations collected prior to 1955 after which the subspecies is thought to have gone extinct. Effective in version 2.0.
  • Cyprinella lutrensis: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella sp 1 (Rio Conchos Shiner) and C. proserpina we must examine specimens and can't make determinations based solely on geography. We maintain determinations of C. lutrensis for records acquired with that name unless we can unambiguously change the name based on geography or have examined the specimen/s. Effective in version 3.0.
  • Cyprinella sp (Nueces River Shiner): We follow Richardson and Gold (1995) and Schönhuth and Mayden (2010) in recognizing this as a distinct form and now refer all records from the Nueces River upstream of Uvalde formerly recognized as Cyprinella lutrensis to this undescribed species. Effective in version 2.0.
  • Cyprinella sp 1 (Rio Conchos Shiner): We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella suavis and Cyprinella panarcys we must examine specimens and can't make determinations based solely on geography.
  • Cyprinella rutila: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Effective in version 3.0.
  • Cyprinella suavis: We follow Schönhuth and Mayden (2010) in recognizing this species and its distribution. However, the American Fisheries Society' common names for fishes publication (Page et al., 2013) does not recognize this taxon at the species level at this time. At the time of this writing we are only recognizing specimens genetically determined by Schönhuth and Mayden (2010) as this species. They note three specimens as this species (UAIC 15313.01, UAIC 8352.02, and MNCN 2433). Those in bold are records not yet included in the Fishes of Texas database as of version 3.0.
  • Cyprinella whipplei: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Since this species co-occurs with the morphologically similar Cyprinella lutrensis we must examine specimens and can't make determinations based solely on geography.  Effective in version 3.0.
  • Cyprinella xanthicara: We follow Schönhuth and Mayden (2010) in recognizing this distinct species and its distribution. Effective in version 3.0.

Dajaus monticola: Durand et al. (2012) determined that Agonostomus monticola should now be Dajaus monticola. Effective in version 3.0.

Dionda: Texas Dionda are mostly morphologically similar and, when range overlap is not indicated, determinations here were mostly done on the basis of ranges defined in Schönhuth et al. (2012). Effective in version 2.0.

  • Dionda sp 1 (Conchos Roundnose Minnow): We follow Schönhuth et al. (2012) in referring all Dionda from the Conchos River (Mex) and into TX (see Figure 1 in that paper) to this undescribed species. Effective in version 2.0.
  • Dionda sp 3 (Colorado Roundnose Minnow): We follow Schönhuth et al. (2012) in referring all Dionda from the Upper Colorado River (TX) (see Figure 1 in that paper) to this undescribed species. Effective in version 2.0.
  • Dionda sp 4 (Upper Pecos Roundnose Minnow): We follow Schönhuth et al. (2012) in referring all Dionda from the Upper Pecos River (TX) (see Figure 1 in that paper) to this undescribed species. Effective in version 2.0.

Eleotris amblyopsis: Texas specimens formerly referred to E. pisonis were determined by Pezold and Cage (2002) to E. amblyopsis. We follow that for all Texas records. Effective in version 3.0.

Erimyzon claviformis: Texas specimens formerly referred to E. oblongus were determined by Bailey et al. (2004) to E. claviformis. We follow that for all Texas records. Effective in version 2.0.

Etheostoma thompsoni: Etheostoma asprigene in the Sabine and Neches Rivers of Texas are now referred to E. thompsoni as per Suttkus et al. (2012). Effective in version 2.0.

Etheostoma pulchellum: Following Bossu and Near (2009) and Bossu et al. (2013), all records of E. spectabile in Texas are now referred to E. pulchellum. Effective in version 2.0.

Etheostoma artesiae: Texas specimens formerly recognized as Etheostoma whipplei are now recognized to be E. artesiae (Piller et al. 2001). Effective in version 1.0.

Fowlerichthys radiosusAntennarius radiosus is now Fowlerichthys radiosus based on genetic analysis by Arnold and Pietsch (2012). This change had been adopted in Fishbase and iNaturalist as well. Effective in version 3.0.

Fundulus blairae: Fundulus notti, F. blairae and F. dispar are all valid, but morphologically very similar species. We follow Fishes of Alabama (Boschung et al.; 2004) and recognize that of these, only F. blairae is found in Texas, and thus specimens from the state formerly referred to F. dispar and F. notti are now called F. blairae. Effective in version 1.0.

Fundulus kansae: F. zebrinus from the Canadian River drainage are now referred to F. kansae as per Kreiser (2001). Effective in version 1.0.

Fundulus xenicus: The genus Adinia was synonymized with Fundulus by Ghedotti and Davis (2013). Effective in version 3.0.

Gambusia krumholzi: G. clarkhubbsi was determined by Echelle et al. (2013) to be a synonym of G. krumholzi. Effective in version 2.0.

Gila: Rio Grande Gila are mostly morphologically similar and determinations here were mostly made on the basis of ranges defined in Schönhuth et al. (2014). Effective in version 3.0.

  • Gila nigrescens: We follow Schönhuth et al. (2014) in referring all Gila from the Guzman drainage (see Figure 1 in that paper) to this species. Effective in version 3.0.
  • Gila pandora: We follow Schönhuth et al. (2014) in referring all Gila from the upper Pecos and Rio Grande drainages (see Figure 1 in that paper) to this species. Effective in version 3.0.
  • Gila pulchra: We follow Schönhuth et al. (2014) in referring all Gila from the Conchos drainage (see Figure 1 in that paper) to this species. Effective in version 3.0.

Gobionellus oceanicus: Gobionellus hastatus and G. gracillimus were synonymized with G. oceanicus by Pezold and Grady (1989) which we adhere to. Effective in version 3.0.

Gymnura lessae: G. micrura was split by Yokata and Carvalho (2017). Those occurring in Texas are now G. lessae. Effective in version 3.0.

Hypanus americanus: Dasyatidae was revised by Last et al. (2016) based on morphological and genetic evidence. Dasyatis americana are now Hypanus americanus. Effective in version 3.0.

Hypanus sabinus: Dasyatidae was revised by Last et al. (2016) based on morphological and genetic evidence. Dasyatis sabina are now Hypanus sabinus. Effective in version 3.0.

Hypanus say: Dasyatidae was revised by Last et al. (2016) based on morphological and genetic evidence. Dasyatis say are now Hypanus say. Effective in version 3.0.

Hypleurochilus multifilis: Hypleurochilus geminatus in Texas were synonymized by Bath (1994) to H. multifilis . That change was accepted by Carpenter (2002) and McEachran and Fechhelm (2010), and is now accepted here. Effective in version 2.0.

Hypostomus sp (Armadillo del Rio): Identifications of Hypostomus specimens from Texas (all introduced), are exceedingly difficult and probably aquarium trade hybrids so we here refer to all as "Hypostomus sp (Armadillo del Rio)". Effective in version 3.0. Previously called "Hypostomus" in earlier versions.

Hyporthodus flavolimbatus: Epinephelus flavolimbatus is now Hyporthodus flavolimbatus according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Hyporthodus mystacinus: Epinephelus mystacinus is now Hyporthodus mystacinus according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Hyporthodus nigritus: Epinephelus nigritus is now Hyporthodus nigritus according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Hyporthodus niveatus: Epinephelus niveatus is now Hyporthodus niveatus according to Smith and Craig (2007) and Craig and Hastings (2007). Effective in version 3.0.

Ictalurus sp (Chihuahua Catfish): Identifications to this undescribed species are difficult, but our research supports validity of this species (Hendrickson, 2017). We here recognize determinations from an unpublished manuscript by Julian Humphries and Robert Rush Miller (1998), specifically UMMZ 214590, 203009, 201537, 203240, 161739, 196733, 66188, TNHC 4240, 2037, UANL 1892, 7121, 2153, 5661, ASU 64-0859, FMNH 100478, MSB 3180, 7778, 6395, 6392. Also see Varela Romero, 2020 and Pérez-Rodríguez, R. 2022, which are both relevant to recognition of the species. Effective in version 2.0. Those in bold are records not yet included in the Fishes of Texas database as of version 3.0.

Ictalurus sp (Rio Grande Blue Catfish): Rodiles-Hernandez et al. (2010), though not indicating the form to be a distinct species, identified morphological differences that support the hypothesis that spotted blue catfishes from the Rio Grande basin are distinctive. Though further study is needed, we here now refer specimens from this drainage known to have vertebral counts (provided by John Lundberg) of 55 to this form. Effective in version 2.0.

Labidesthes vanhyningi: We now recognize this species that is sympatric and morphologically similar to L. sicculus. Werneke and Armbruster (2015) based on morphological data, determined the species to exist in the Neches, Sabine and Red River drainages of Texas. Specimens of L. sicculus in east Texas now need to be examined to determine the range of the species. We have re-determined records indicated by the authors as this species from our study area (TU106407, TU200888, TU200889, NCSM36316, NCSM36612, NCSM37403, NCSM37416, NCSM55222, UAIC778.02, UT158.22, UT158.41, AUM26380, TU114924, UF_29501).  Those in bold are records not yet included in the Fishes of Texas database as of version 3.0.

Lepomis aquilensis: We recognize all specimens in our study area previously identified as Lepomis megalotis as L. aquilensis as per genetic and morphological work by Kim et al. (2021).

Lepomis miniatus: We now recognize synonymy by Warren (1992) of Lepomis punctatus in Texas to L. miniatus. Effective in version 1.0.

Luxilus chrysocephalus: Based on biochemical analysis by Dowling and Moore (1984) we recognize specimens in our study area previously identified as L. cornutus as L. chrysocephalus.

Macrhybopsis: Texas species in this genus are morphologically similar and, when range overlap is not indicated, determinations here are now based mostly on the ranges defined by Eisenhour (2004). Effective in version 1.0.

Menidia beryllina: Morphologically similar to M. beryllina, M. audens was recently accepted by the American Fisheries Society (Page et al. 2013) and others. Morphological evidence provided by Suttkus et al. (2005) supports taxonomic separation from M. beryllina, however genetic support by Fluker et al. (2011) is weaker, but suggests some differentiation. Effective in version 1.0, 2.0, and 3.0.

Micropterus nigricans(change not yet applied): We accept a revision, based on genetic evidence (Kim et al., 2022), which indicates that, within our study area, the name M. nigricans should apply to what was previously called M. salmoides. We also now lump a subspecies we once recognized, M. salmoides nuecensis, within M. nigricans as there is now little genetic evidence suggesting it as a subspecies. Effective in version 3.0. (note: Prior to Kim et al (2022) we followed Lutz-Carrillo et al. (2006) in recognizing this form. Based on their genetic analysis, the form apparently persists to some degree in the Devils River as genetically unique lineage. We never had sufficient morphological characters to differentiate the subspecies from other M. salmoides (Now M. nigricans) and had in versions 1.0 to 2.0 identified records of M. salmoides collected 1940 or before from the Nueces and Rio Grande as this sub species.) 

Micropterus salmoides(change not yet applied): We accept a revision, based on genetic evidence (Kim et al., 2022), which indicates the name M. salmoides should apply to what was previously called the Florida Bass, M. floridanus. (Note: Records within our study area that were previously determined as M. salmoides are now M. nigricans (see above).)  Effective in version 3.0.

Notropis megalops: Notropis amabilis from parts of the Rio Grande drainage have been genetically determined to be N. megalops by Conway and Kim (2016). We adjust all N. amabilis from the lower Pecos River and its tributaries to N. megalops - this is where N. megalops does not co-occur with N. amabilis. But, since N. amabilis and N. megalops are sympatric in the Devils River and Sycamore Creeks (and elsewhere) we cannot make geographic-based determinations within the entire range of N. megalops. We applied N. megalops determinations for those specific occurrences where supplied by Conway and Kim (2016), listed here: TCWC 11890.01, TCWC 15558.12, TCWC 16323.15, TCWC 16456.11, TCWC 16457.08, TCWC 6242.03, TCWC 6243.05, TCWC 6669.09, TCWC 7510.11, USNM 427772, USNM 70, MCZ 1682, MCZ 171862, OMNH 65168, TCWC 7515.08, TCWC 11044.01, TCWC 1159.01, TCWC 11880.01, TCWC 11882.01, TCWC 11884.01, TCWC 11885.01, TCWC 11888.01, TCWC 11888.11, TCWC 11889.01, TCWC 13084.10, TCWC 13085.06, TCWC 13087.08, TCWC 13096.10, TCWC 14774.01, TCWC 14781.01, TCWC 15558.02, TCWC 16323.14, TCWC 16325.16, TCWC 16455.05, TCWC 16456.10, TCWC 16457.02, TCWC 3907.07, TCWC 6241.01, TCWC 6669.06, TCWC 7509.07, TCWC 7510.09, TCWC 7513.03, TCWC 7514.03, TCWC 7515.02, TCWC 7516.06, TCWC 7517.04, TCWC 7518.04, TCWC 7519.04, TCWC 7523.07, TCWC 7528.02, TCWC 7529.04, TCWC 7532.04, TCWC 7533.08, TNHC 15972, TNHC 16027, TNHC 27499, TNHC 29455, TNHC 58718, TNHC 9461, TU 5945, UAIC 09882.05, USNM 129, USNM 39654. Effective in version 3.0.

Notropis simus pecosensis: We recognize this subspecies described by Chernoff et al. (1982). Effective in version 2.0.

Notropis simus simus: We recognize this subspecies described by Chernoff et al. (1982). Effective in version 2.0.

Notropis suttkusi: All records determined as N. percobromis or N. rubellus are called N. sutkussi as per Humphries and Cashner (1994). A map of the species in the N. rubellus complex is provided by Berendzen et al (2008).

Percina apristis: We follow Robins and Page (2007) in referring what was formerly called Percina sciera from the Guadalupe drainage to P. apristis. Effective in version 1.0.

Pterygoplichthys sp (Sailfin Catfish): At least 3 species of Pterygoplichthys have been introduced in Texas (P. anisitsi, P. disjunctivus and P. multiradiatus) and are likely hybridizing, as discussed by Hoover et al. (2004). Due to the taxonomic confusion, we are recognizing them under a single name. Effective in version 3.0.

Syngnathus scovelli: Syngnathus affinis is no longer recognized and is now a synonym of S. scovelli (Tolan 2008). Effective in version 2.0.

Incoming literature that will influence future taxonomic decisions:

Cyprinidae: Susana Schönhuth, Jasna Vukić, Radek Šanda, Lei Yang, Richard L. Mayden, Phylogenetic relationships and classification of the Holarctic family Leuciscidae (Cypriniformes: Cyprinoidei), Molecular Phylogenetics and Evolution, Volume 127, 2018, Pages 781-799, ISSN 1055-7903, https://doi.org/10.1016/j.ympev.2018.06.026.

Dorosoma petenenseElías, D. J., McMahan, C. D., & Piller, K. R. (2021). Molecular data elucidate cryptic diversity within the widespread Threadfin Shad (Dorosoma petenense: Clupeidae) across the Nearctic and Northern Neotropics. Hydrobiologia. https://doi.org/10.1007/s10750-021-04713-8

Etheostoma lepidumMacGuigan, D. J., Hoagstrom, C. W., Domisch, S., Hulsey, C. D., & Near, T. J. (2021). Integrative ichthyological species delimitation in the Greenthroat Darter complex (Percidae: Etheostomatinae). Zoologica Scripta, 00, 1 27. https://doi.org/10.1111/zsc.12504

Cyprinus carpio (recognition of C. rubrofuscus)

  • Zhou, J., Wu, Q., Wang, Z. and Ye, Y. (2004). Molecular Phylogenetics of Three Subspecies of Common carp Cyprinus Carpio, based on sequence analysis of cytochrome b and control region of mtDNA. Journal of Zoological Systematics and Evolutionary Research 42(4): 266–269.
  • Xu, P., Zhang, X., Wang, X., et al. (2014). Genome sequence and genetic diversity of the common carp, Cyprinus carpio. Nature Genetics 46: 1212–1219 doi:10.1038/ng.3098

Sander: https://afspubs.onlinelibrary.wiley.com/doi/full/10.1002/fsh.10582

Amia calva: https://royalsocietypublishing.org/doi/pdf/10.1098/rsbl.2022.0395

Hemichromis guttatus: https://link.springer.com/article/10.1007/s10750-022-05060-y

Literature 

Arnold, R. J., and T. W. Pietsch. 2012. Evolutionary history of frogfishes (Teleostei: Lophiiformes: Antennariidae): a molecular approach. Molecular Phylogenetics and Evolution 62:117-129. [Available online 2 Oct. 2011; publisher listed hard copy as 2012].

Bailey, R. M., W. C. Latta, and G. R. Smith. 2004. An Atlas of Michigan Fishes with Keys and Illustrations for Their Identification. http://deepblue.lib.umich.edu/handle/2027.42/56435.

Bath, H. 1994. Untersuchung der arten Hypleurochilus geminatus (Wood 1825), H. fissicornis (Quoy & Gaimard 1824) und H. aequipinnis (Gunther 1861), mit revalidation von Hypleurochilus multifilis (Girard 1858) und beschreibung von zwei neuen arten (Pisces: Blenniidae). Senckenbergiana Biologica 74:59-85.

Berendzen, P. B., A. M. Simons, R. M. Wood, T. E. Dowling, and C. L. Secor. 2008. Recovering cryptic diversity and ancient drainage patterns in eastern North America: historical biogeography of the Notropis rubellus species group (Teleostei: Cypriniformes). Mol. Phylogenet. Evol. 46:721737.  doi:10.1016/j.ympev.2007.07.008

Bessert, M. 2006. Molecular Systematics and Population Structure in the North American Endemic Fish Genus Cycleptus (Teleostei: Catostomidae). Dissertations and Theses in Biological Sciences, December. http://digitalcommons.unl.edu/bioscidiss/1.

Boschung, H. T., R. L. Mayden, and J. R. Tomelleri. 2004. Fishes of Alabama. Washington, D.C: Smithsonian Books.

Bossu, C. M., and T. J. Near. 2009. Gene trees reveal repeated instances of mitochondrial DNA introgression in Orangethroat Darters (Percidae: Etheostoma). Systematic Biology 58 (1):114-29. doi:10.1093/sysbio/syp014.

Bossu, C. M., J. M. Beaulieu, P. A. Ceas, and T. J. Near. 2013. Explicit tests of palaeodrainage connections of southeastern North America and the historical biogeography of Orangethroat Darters (Percidae: Etheostoma: Ceasia). Molecular Ecology 22: 5397-5417.

Brown, W. H. 1953. Introduced fish species in the Guadalupe River basin. Texas Journal of Science 5:245-251.

Burr, B. M., and R. L. Mayden. 1999. A new species of Cycleptus (Cypriniformes: Catostomidae) from Gulf Slope drainages of Alabama, Mississippi, and Louisiana, with a review of the distribution, biology and conservation status of the genus. Bulletin of the Alabama Museum of Natural History 20:20-58.

Buth, D. G., R. L. Mayden, and J. D. McEachran. 2001. Allozymic and isozymic evidence for polytypy in the North American catostomid genus Cycleptus. Copeia 2001:899-906. doi:10.1643/0045-8511(2001)001[0899:AAIEFP]2.0.CO;2.

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