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Verlag Dr. Friedrich PfeilISSN 0936-9902

Ichthyological Exploration of Freshwaters

Volume 21Number 1

An international journal for fi eld-orientated ichthyology

Ichthyological Exploration of FreshwatersAn international journal for fi eld-orientated ichthyology

Volume 21 • Number 1 • March 2010pages 1-96, 62 fi gs., 14 tabs.

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Ichthyol. Explor. Freshwaters, Vol. 21, No. 1

Ichthyol. Explor. Freshwaters, Vol. 21, No. 1, pp. 63-72, 3 figs., 1 tab., March 2010© 2010 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902

An overview of North America’s diminutive freshwater fish fauna

Micah G. Bennett* and Kevin W. Conway*

Based on a review of the literature, we compile a table of North American freshwater fishes with maximum standard length (SL) less than or equal to 50 mm and information on size at sexual maturity. Using previously published criteria for miniature fishes (< 26 mm maximum SL, or those that mature at SL < 20 mm), we found evidence for seven miniature fishes on the North American continent: Elassoma alabamae, E. zonatum, Leptolucania ommata, Heterandria formosa, Megupsilon aporus, Cyprinodon macularius and Notropis saladonis, the smallest being E. alabamae (Spring Pygmy Sunfish, maximum size 25.5 mm SL). We compare the number of miniature species in North America to those from Africa, South Asia and South America and discuss patterns of distribution, habitat preference and imperilment amongst North America’s smallest freshwater fishes.

* Department of Biology, Saint Louis University, 3507 Laclede Avenue, St. Louis, MO 63103, USA. E-mail: [email protected], [email protected]

Introduction

The North American continent contains one of the most diverse temperate freshwater fish faunas in the World (Abell et al., 2000; Lundberg et al., 2000), with over 1050 described and numerous undescribed species (Page & Burr, 1991; Lundberg et al., 2000). At the turn of the 20th century the North American ichthyofauna was credited with the distinction of having some of the “smallest known fishes and at the same time the smallest known vertebrates” (Smith, 1902). In more recent years the North American continent has lost its “smallest vertebrate” status, which has passed back and forth between Indo-Pacific, Australa-sian and South East Asian countries (Herre, 1929; Winterbottom & Emery, 1981; Watson & Walker, 2004). Among these is the recent description of the cypriniform genus Paedocypris (Kottelat et al., 2006), which contains the smallest living verte-

brate species (P. progenetica), maturing at sizes under 8 mm in standard length (SL). Though small-bodied fishes have always been of interest, ichthyologists have only fairly re-cently began to undertake continent-wide assess-ments of small-bodied species. Weitzman & Vari (1988) undertook the first such survey document-ing the numerous miniature freshwater fish spe-cies of South America. They defined miniature fishes as those no greater than 26 mm SL or ma-turing at sizes smaller than 20 mm SL and identi-fied 85 such species. Costa & Le Bail (1999) added 26 more to the count for South America. Using Weitzman & Vari’s criterion, Kottelat & Vidthayanon (1993) provided a list of miniature fishes for south and southeast Asia and, more recently, Conway & Moritz (2006) compiled a similar list for Africa. Besides providing a more thorough knowledge of continental biodiversity, identifying and tabulating the number of small-

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bodied fishes is important to largescale ecological and evolutionary investigations such as those focused on geographic gradients in body size (e.g., Belk & Houston, 2002; Knouft, 2004), phy-logenetic trends in body size (e. g., Knouft & Page, 2003; Ruber et al., 2007), life history evolution (e. g., Weitzman & Vari, 1988), and issues of con-servation concern, such as size-biased extinction risks (e. g., Olden et al., 2007). In this paper we provide a list of all North American freshwater fishes with a maximum SL less than or equal to 50 mm in order to assess whether the North American ichthyofauna con-tains any miniature species of fish (sensu Weitz-man & Vari, 1988) and to determine the smallest North American fish species.

Methods

We conducted surveys of peer-reviewed literature and books on fishes and fish biology using four standard publication databases (Web of Science, Biological Abstracts, Google Scholar, JSTOR) and compiled a table of all freshwater fishes with a maximum standard length (SL) of less than or equal to 50 mm that inhabit North America (in-cluding Mexico). This was necessarily an arbitrary cut-off point, but it was chosen in order to include the lower tail of the size distribution of North American freshwater fishes (sensu Knouft & Page, 2003) and to include any miniature species (< 26 mm maximum SL, sensu Weitzman & Vari, 1988). We used the University of Michigan Mu-seum of Zoology online catalogue of the Division of Fishes (UMMZ, 2009) which includes SL infor-mation to determine if specimens larger than those documented in the literature survey had been collected for species for which maximum standard lengths had been published. That catalogue also served as a source of information for species for which no literature could be found. Literature information was also used to compile data on the minimum SL at maturity of all species with standard lengths of less than 50 mm in order to assess whether any species included in our survey matured at sizes below 20 mm SL. Lastly, we utilized the most recent status rankings of the American Fisheries Society (Jelks et al., 2008) to examine patterns of imperilment among diminu-tive fishes.

Results

Our survey documented 146 species of North American freshwater fishes with a maximum SL less than, or equal to, 50 mm, 6 of which mature at sizes of less than 20 mm SL (Table 1). The pool of diminutive fishes (equal to or below 50 mm SL) covers a broad range of North American fish families but is made up largely of Poeciliidae (32.0 %), Cyprinidae (18.4 %), Cyprinodontidae (17.0 %), and Percidae (14.3 %). Three of these, the Cyprinidae, Percidae and Poeciliidae are the most diverse families of fishes in North America, making up about 26 %, 16 % and 8 %, respec-tively, of that fauna (Page & Burr, 1991; Lundberg et al., 2000; Jelks et al., 2008), whereas the Cyprin-odontidae accounts for a much smaller portion of that fauna (less than 4 %; Jelks et al., 2008). Thus the composition of Table 1 is not merely reflective of the overall pattern of North American freshwater fish diversity. Using the definition of Weitzman & Vari (1988) seven of these 146 species are considered miniature: Elassoma alabamae (Fig. 1), Leptolucania ommata (Fig. 2), Heterandria formosa, Megupsilon aporus (Fig. 3), Cyprinodon macularius, Notropis saladonis and Elassoma zon-tatum. Elassoma alabamae is the only species con-sidered a miniature based on maximum known size (25.5 mm SL). The remaining miniatures are defined as such based on minimum size at ma-turity as all achieve maximum standard lengths greater than 26 mm.

Discussion

In comparison to other continents for which lists of miniature freshwater fishes have been compiled (South America – Weitzman & Vari, 1988; South-east Asia – Kottelat & Vidthayanon, 1993; Africa – Conway & Moritz, 2006) North America has few truly miniature freshwater fish species. Af-rica contains 24 miniature freshwater fish species (Conway & Moritz, 2006), South America harbors well over 100 such species (Weitzman & Vari, 1988; Costa & Le Bail, 1999), and Southeast Asia contains over 50 species (Kottelat & Vidthayanon, 1993; Kottelat et al., 2006), compared to only seven species in North America (Table 1). There is currently no published list of European mini-ature species but based on Kottelat & Freyhof (2007) there appears to be just two: Economidich-thys trichonis (maximum size 25 mm SL) and

Bennett & Conway: Diminutive fishes of North America

65


a b

Fig. 1. Elassoma alabamae; U.S.A.: Alabama: Limestone County: Beaverdam Creek. a, male; b, female. Photographs courtesy of P. Rakes and J. R. Shute.

Fig. 2. Leptolucania ommata; locality unknown. a, male. b, female. Photographs courtesy of P. Loiselle.

a

b

66

Table 1. North American freshwater fishes with a maximum known standard length (SL) less than or equal to 50.0 mm. All measurements are in SL unless otherwise stated. Taxa highlighted in bold are miniature species (sensu Weitzman and Vari 1988). Conservation status abbreviations (following Jelks et al. 2008): E, endangered; T, threatened; V, vulnerable; X, extinct; Xn, extirpated in nature; Xp, possibly extinct; –, not assessed.

order – family/speciesmax SL(mm)

min SL at maturity

(mm)

reference (maximum known SL;minimum known SL at maturity) C

onse

rvat

ion

Stat

us

Cypriniformes – CyprinidaeCyprinella alvarezdelvillari 43.8 25.0 (M) Miller et al., 2005: 110 EDionda diaboli 48 ? Miller et al., 2005: 116 EDionda erimyzonops 39 21.0 (M) Miller et al., 2005: 117; Hubbs & Miller, 1974 –Evarra tlahuacensis 48 ? Miller et al., 2005: 120 XExtrarius n. sp. cf. aestivalis 50 ? Gilbert, 1992 rare?Hybopsis amecae 41 ? Miller et al., 2005:127 –Hybopsis aulidion 39 ? Miller et al., 2005: 130 –Iotichthys phlegethontis 41 ? Miller & Behnke, 1985 ELythrurus alegnotus 50 ? Boschung & Mayden, 2004 –Lythrurus snelsoni 43.2 ? Robison, 1985 VNotropis alborus 50 ? Hubbs & Raney, 1947: 11 –Notropis ammophilus ‘ca.’ 48.0 29.0 (M); 31.0 (W) Boschung & Mayden, 2004 –Notropis anogenus 48 ? Bailey, 1959 TNotropis buchanani 41 ? Miller et al., 2005: 132 –Notropis calbazas 26.8 ? Lyons & Mercado-Silva, 2004 ENotropis chalybaeus 50 ? Boschung & Mayden, 2004 VNotropis harperi 49 25.0 (W) Boschung & Mayden, 2004 –Notropis melanostomus 38 21 Boschung & Mayden, 2005; Suttkus & Bailey, 1990 TNotropis ortenburgeri 50 ? UMMZ, 2009 VNotropis rafinesquei ‘ca.’ 43.0 22.5 Haag et al., 2007 –Notropis sabinae 49 38.0 (W; TL) Heins, 1981; Williams & Bonner, 2006 –Notropis saladonis 38 19.0 (M); 21.0 (W) Miller et al., 2005: 135; Hubbs & Hubbs, 1958 XpNotropis tropicus 41 27.0 (M) Miller et al., 2005:137 –Oregonichthys kalawatseti 49.2 ? Markle et al., 1991: 280 VStypodon signifer 40 ? Miller et al., 2005: 140 XLythrurus roseipinnis ‘ca.’ 50.0 28 Boschung & Mayden, 2004 –Characiformes – CharacidaeHyphessobrycon compresus 46 ? Miller et al., 2005: 158 –Siluriformes – IctaluridaeNoturus stanauli 36.2 ? Etnier & Jenkins, 1980 EPercopsiformes – AmblyopsidaeAmblyopsis rosae 43 ? UMMZ, 2009 TChologaster cornuta 48 20.0 (W) Ross & Rhode, 2003: 110 –Atheriniformes – AtherinopsidaeAtherinella lisa 45 ? Miller et al., 2005: 189 ECyprinodontiformes – CyprinodontidaeCualac tessellatus 45.5 ? Miller et al., 1956: 9 ECyprinodon arcuatus 42 ? Minckley et al., 2003 XpCyprinodon atrorus 35 ? Miller et al., 2005: 298 ECyprinodon beltrani 41 ? Miller et al., 2005: 299 VCyprinodon diabolis 28 ? UMMZ, 2009 ECyprinodon elegans 49 ? Suttkus & Williams, 2008 ECyprinodon eremus 41.5 ? Minckley et al., 2005 ECyprinodon eximius 46 ? Miller et al., 2005: 303 TCyprinodon fontinalis 50 ? Miller et al., 2005: 303 ECyprinodon labiosus 40 ? Miller et al., 2005: 304 E


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Cyprinodon latifasciatus 43 ? Miller et al., 2005: 304 XCyprinodon macrolepis 44 ? Miller et al., 2005: 305 ECyprinodon macularius 37.8 15 Minckley et al., 2004; Schoenherr, 1988 ECyprinodon meeki 49 ? Miller et al., 2005: 307 ECyprinodon nazas 46 ? Miller et al., 2005: 308 TCyprinodon pecosensis ‘ca.’ 50.0 ? Garrett, 1982 ECyprinodon pisteri 49.3 23.0-25.0 (M) Minckley et al., 2002: 688, 689 ECyprinodon rubrofluviatilis 48 ? UMMZ, 2009 –Cyprinodon simus 33 ? Miller et al., 2005: 309 ECyprinodon suavium 45.5 ? Strecker, 2003 –Cyprinodon variegatus hubbsi 35 ? Gilbert, 1992 VCyprinodon verecundus 35 ? Humphries, 1984: 63 EGarmanella pulchra 38 ? Miller et al., 2005: 315 –Jordanella floridae 39 ? McEachran & Fechhelm, 1998:899 –Megupsilon aporus 36 15.0 (M); 18.0 (W) Miller et al., 2005: 317 XnCyprinodontiformes – FundulidaeAdenia xenica 42 ? Boschung & Mayden, 2004 –Leptolucania ommata 27 12.0 (M); 15.0 (W) Boschung & Mayden, 2004 –Lucania goodei 42 ? Boschung & Mayden, 2004 –Lucania interioris ~35.0 ? Miller et al., 2005: 316 ECyprinodontiformes – GoodeidaeAllodontichthys polylepis 50 ? Miller et al., 2005: 269 EAllotoca maculata 48 ? Miller et al., 2005: 274 EAllotoca meeki 50 ? Miller et al., 2005: 274 EAllotoca zacapuensis 50 ? Miller et al., 2005: 275 ECharacodon garmani 27 ? Miller et al., 2005: 279 XCrenichthys nevadae 44 ? Hubbs, 1932 TCyprinodon pachycephalus 50 ? Miller et al., 2005: 308 ESkiffia bilineata 42 ? Miller et al., 2005: 284 ESkiffia francesae 43 ? Miller et al., 2005: 284 XnCyprinodontiformes – PoeciliidaeBrachyrhaphis hartwegi 43 ? Miller et al., 2005: 219 –Gambusia alvarezi 38 ? Miller et al., 2005: 221 EGambusia atrora 40 ? Miller et al., 2005: 221 –Gambusia aurata 34 ? Miller et al., 2005: 222 –Gambusia clarkhubbsi 43 ? Garrett & Edwards, 2003 EGambusia eurystoma 36 ? Miller et al., 2005: 223 VGambusia hurtadoi 36 ? Miller et al., 2005: 223 EGambusia krumholzi 38 ? Miller et al., 2005: 223 VGambusia longispinis 29 ? Miller et al., 2005: 224 EGambusia regani 42 32.0 (W) Miller et al., 2005: 225 –Gambusia senilis 46 ? Miller et al., 2005: 226 TGambusia sexradiata 43 ? Miller et al., 2005: 227 –Gambusia speciosa 48 ? Miller et al., 2005: 227 TGambusia vittata 37 25.0 (W) Miller et al., 2005: 228 –Heterandria formosa 30 15.0 (W) Boschung & Mayden, 2004 –Heterophallus echeagarayi 35 ? Miller et al., 2005: 230 –Heterophallus milleri 31 ? Miller et al., 2005: 231 –Heterophallus rachovii 32 ? Miller et al., 2005: 231 –Phallichthys fairweatheri 46 ? Miller et al., 2005: 232 –Poecilia chica 46 ? Miller et al., 2005: 234 VPoecilia sulphuraria 50 ? Miller et al., 2005: 239 TPoeciliopsis baenschi 44 ? Miller et al., 2005: 241 –Poeciliopsis balsas 43 ? Miller et al., 2005: 242 –Poeciliopsis fasciata 45 ? Miller et al., 2005: 242 –Poeciliopsis latidens 48 ? Miller et al., 2005: 245 T

Table 1. (continued).

68

Poeciliopsis lucida ~42.0 ? Miller et al., 2005: 246 –Poeciliopsis monacha 44 ? Miller et al., 2005: 246 –Poeciliopsis occidentalis 48 ? Miller et al., 2005: 246 EPoeciliopsis presidionis ~45.0 ? Miller et al., 2005: 248 –Poeciliopsis prolifica 42 ? Miller et al., 2005: 248 –Poeciliopsis viriosa 46 ? Miller et al., 2005: 251 –Priapella bonita 45 ? Miller et al., 2005: 251 XPriapella intermedia 45 ? Miller et al., 2005: 253 –Xiphophorus andersi 40 ? Miller et al., 2005: 255 –Xiphophorus clemenciae 46 ? Miller et al., 2005: 255 TXiphophorus continens ~30.0 ? Miller et al., 2005: 256 –Xiphophorus cortezi 50 ? Miller et al., 2005: 257 –Xiphophorus couchianus 37 ? Miller et al., 2005: 257 EXiphophorus evelynae 48 ? Miller et al., 2005: 257 –Xiphophorus gordoni 30 ? Miller et al., 2005: 258 EXiphophorus maculatus 45 ? Miller et al., 2005: 259 –Xiphophorus meyeri 46.2 ? Miller et al., 2005: 260 EXiphophorus milleri 35 ? Miller et al., 2005: 261 EXiphophorus multilineatus ~45.0 ? Miller et al., 2005: 262 –Xiphophorus nigrensis 42 ? Miller et al., 2005: 263 –Xiphophorus pygmaeus 37.2 ? Morris & Ryan, 1995 –Xiphophorus xiphidium 49 ? Miller et al., 2005: 264 –Cyprinodontiformes – RivulidaeMillerichthys robustus 33 ? Miller et al., 2005: 207 ERivulus tenuis 40 ? Miller et al., 2005: 208 –Perciformes – ElassomatidaeElassoma alabamae 25.5 ? Mayden, 1993 EElassoma boehlkei 27 ? Rohde & Arndt, 1987 TElassoma evergladei 28 ? Boschung & Mayden, 2004 –Elassoma okatie 28.7 ? Rohde & Arndt, 1987 VElassoma okefenokee 28 ? Boschung & Mayden, 2004 –Elassoma zonatum 38.5 19.2 (W) Walsh & Burr, 1984: 36, 41Perciformes – PercidaeEtheostoma australe 50 ? Miller et al., 2005: 335 EEtheostoma chlorosoma 50 ? Page, 1983 –Etheostoma denoncourti 28.6 ? Stauffer & Van Sink, 1997 VEtheostoma ditrema 45 ? Page, 1983 TEtheostoma edwini 49 27.0 (W) Boschung & Mayden, 2004 –Etheostoma fonticola 35.5 24.4 (W TL) Burr, 1978; Schenk & Whiteside, 1977 EEtheostoma fusiforme 46 32.0 (TL) Boschung & Mayden, 2004 –Etheostoma gracile 49 35.0 (TL) Boschung & Mayden, 2004 –Etheostoma grahami 44 ? Miller et al., 2005: 336 TEtheostoma lugoi 37.6 ? Norris & Minckley, 1997: 162 EEtheostoma microperca 37 ~28.0 Etnier & Starnes, 1993 –Etheostoma nuchale 45 ? Boschung & Mayden, 2004 EEtheostoma okaloosae 48 ? Burkhead et al., 1992 EEtheostoma pottsi 50 ? Miller et al., 2005: 337 TEtheostoma proeliare 40 29.2 (M); 30.7 (W) Boschung & Mayden, 2004 –Etheostoma rubrum 46 27.0 (W) Raney & Suttkus, 1966; Knight & Ross, 1992:102 EEtheostoma segrex 43.8 ? Norris & Minckley, 1997: 171 EEtheostoma stigmaeum 48.9 35.6 (M); 35.7 (W) Boschung & Mayden, 2004 –Etheostoma tippecanoe 36 22.5 (M); 21.2 (W) Stauffer & Van Sink, 1997; Warren et al., 1986 VEtheostoma trisella 49 37 Boschung & Mayden, 2004; Ryon, 1986: 81 EEtheostoma zonifer 38 ? Boschung & Mayden, 2004 –Scorpaeniformes – CottidaeCottus paulus 38 25 Williams, 2002; Boschung & Mayden, 2004 E

Table 1. (continued).


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Knipowitschia goerneri (maximum size 22 mm SL). Any explanation for this disparity is at this time highly speculative; however, Africa, Asia and South America differ greatly from North America in several key aspects that may promote miniaturization and diversification of diminutive fishes. The equatorial position of much of Africa, South and Southeast Asia and South America is likely a major factor in diversification on those continents in general (Lundberg et al., 2000). The large number of species in the tropics has also been suggested to make miniature species simply more likely to evolve (Weitzman & Vari, 1988). Collette (1962) suggested a link between the acidic environments inhabited by Etheostoma fusiforme (Swamp Darter) and its reduced lateral line system and overall small body size. Simi-larly Weitzman & Vari (1988) noted that many South American miniatures occur in this environ-ment, which is common across many portions of the continent. They also recognized that most of the miniature freshwater fishes included in their table, if not found explicitly in acidic waters, in-habited still or slow-flowing habitats or micro-habitats. Kottelat & Vidthayanon (1993) and Kottelat et al. (2006) also commented that most miniature Southeast Asian freshwater fishes oc-cupy lentic or slow-flowing habitats. Our review further supports the notion that miniature fishes are more likely to occur in habitat with little flow. Cyprin odon macularius, Elassoma alabamae and Megupsilon aporus occur in springs or spring-fed pools with little or no flow (Hendrickson & Romero, 1989; Mayden, 1993; Miller & Walters, 1972). Leptolucania ommata and Elassoma zonatum are found in vegetated swamps and backwaters or streams with little flow (Boschung & Mayden, 2004). Heterandria formosa prefers pools of slug-gish, heavily-vegetated streams (Boschung & Mayden, 2004) and Notropis saladonis inhabits slow-flowing, silty streams (Hubbs & Hubbs, 1958). In addition, many of the smallest species in Table 1, while not miniature by the definition of Weitzman & Vari (1993), are found mostly in swamp and spring habitats (e. g. Elassoma, Xipho-phorus). These habitats, as well as smaller streams (even swift-flowing), likely limit dispersal, and thus gene flow, among populations of small-bodied fishes, promoting diversification (Knouft & Page, 2003). While this general model encompasses most of the diminutive species we examined (and in-deed those studied on other continents), there is

Fig. 3. Megupsilon aporus, Mexico: Matinal de El Potosi Nuevo Leon. Male. Photograph courtesy of P. Loi-selle.

also a small component of diminutive North American freshwater fish diversity that inhabits fast flowing, large and medium-sized upland rivers and streams with gravel, cobble, boulder and bedrock as predominant sediments (e. g. Noturus stanauli, Etheostoma tippecanoe, E. denon-courti). Similar habitat preferences have been reported for a small number of miniature taxa from other continents, including the catfishes Hoplomyzon papillatus (Stewart, 1985), Horiomyzon retropinnatus (Stewart, 1986), and the cichlid Nan-nochromis minor (Roberts & Stewart, 1976). The microhabitats associated with the cryptobenthic behavior of these latter species may be character-ized by reduced flow and analogous to the slow velocities of oxbows, springs and swamp. Imperilment was high among diminutive fishes, with 58 % of those listed in Table 1 labeled as vulnerable (9 % of total), threatened (10.2 %), endangered (32.0 %) or extinct (6.8 %) according to Jelks et al. (2008). Only barely over 42 % of these fishes did not warrant listing by Jelks et al. (2008) (i. e. not threatened). Elassoma alabamae is particularly threatened and in need of further protection (Conway & Mayden, 2006). Our review also highlights the lack of critical life history in-formation (minimum size at maturity) for many of the fish species we examined. We anticipate that with more study of reproductive biology, additional species on our list will be identified as miniature based on size at maturity (less than 20 mm SL) in the future. Small bodied freshwater fishes are particu-larly faced with a great threat of extinction, but may get less attention than their larger commer-cially-harvested relatives (Olden et al., 2007). Without a better knowledge of the processes and conditions that gave rise to and help maintain the

70

diversity of small bodied freshwater fishes, and the habitats that they rely on, there will be little hope for preserving these interesting and unique components of global biodiversity.

Acknowledgements

We are grateful to C. Dillman, J. Knouft, N. Lang, R. Mayden and L. Page for discussions on small-bodied North American freshwater fishes, P. Rakes and J. R. Shute (Conservation Fisheries Inc.) for the photographs of Elassoma alabamae, P. Loiselle for the photographs of Leptolucania and Megupsilon and M. Kottelat and R. Vari for helpful comments on an earlier draft of the manu-script. Finally, we thank R. Mayden and R. Wood for their continuing support and advice. This research was supported by Saint Louis University and the National Science Foundation (DEB 0808446 to KWC).

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Received 30 May 2009Revised 29 October 2009

Accepted 3 November 2009


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Articles appearing in this journal are indexed in:

AQUATIC SCIENCES and FISHERIES ABSTRACTSBIOLIS - BIOLOGISCHE LITERATUR INFORMATION SENCKENBERG

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FISHLITZOOLOGICAL RECORD

C O N T E N T S

Kawase, Seigo and Kazumi Hosoya: Biwia yodoensis, a new species from the Lake Biwa/Yodo River Basin, Japan (Teleostei: Cyprinidae) ................................................................... 1

Moritz, Timo: Fishes of Iguidi River – a small forest stream in South-East Benin ................. 9

Silva, Anjana, Kalana Maduwage and Rohan Pethiyagoda: A review of the genus Rasbora in Sri Lanka, with description of two new species (Teleostei: Cyprinidae) ...................... 27

Hu, Yu-Ting and E Zhang: Homatula pycnolepis, a new species of nemacheiline loach from the upper Mekong drainage, South China (Teleostei: Balitoridae) .................................... 51

Bennett, Micah G. and Kevin W. Conway: An overview of North America’s diminutive freshwater fish fauna.................................................................................................................. 63

Costa, Wilson J. E. M.: Simpsonichthys harmonicus, a new seasonal killifish from the São Francisco River basin, northeastern Brazil (Cyprinodontiformes: Rivulidae) .................. 73

Costa, Wilson J. E. M.: Two new species of the Rivulus urophthalmus group from the Tocan-tins and Xingu river drainages, eastern Brazilian Amazon (Cyprinodontiformes: Rivul-idae) .............................................................................................................................................. 79

Román-Valencia, César, Carlos A. García-Alzate, Raquel I. Ruiz-C. and Donald C. Tap-horn B.: A new species of Creagrutus from the Güejar River, Orinoco Basin, Colombia (Characiformes: Characidae) ..................................................................................................... 87

New books – Books received ........................................................................................................... 96


Volume 21 • Number 1 • March 2010

Cover photograph:Biwia yodoensis

Seigo Kawase and Kazumi Hosoya(this volume pp. 1-7)

verlag dr. friedrich pfeil issn 0936-9902 ichthyological...

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