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Rhinichthys gabrielino, new species, Santa Ana Speckled Dace Figs. 3, 4a, 5 Synonymy. Agosia nubila carringtoni Jordan and Evermann 1896:312; Culver and Hubbs 1917:82; Rhinichthys osculus carringtoni Shapovalov and Dill 1950:386; Shapovalov, Dill, and Cordone 1959; Kimsey and Fisk 1960:469; Cornelius 1969:1; Oakey et al. 2004:212; R. osculus subsp. widely used in California since the 1950s (for example: Moyle 2002, Moyle et al. 2015). Holotype. WFB 3498 (Fig. 4a), 73 mm SL, East Fork San Gabriel River, South of Upper Monroe Road, Angeles National Forest, Los Angeles County, California, 34 14’ 15.85”N 117 49’ 14.58”W, USA, 2 July 2021, J. Pareti (California Department of Fish and Wildlife). Paratypes. WFB 3499–3500, WFB 3501–3507 (n =9). Same data as holotype. Meristics: holotype (paratypes): Lengths(mm): standard 73(45–78), fork 82 (54–84), total 89 (57–90) Lateral line scales: 73(69–75). Lateral line complete on all fish, although there were 1–3 scale rows beyond the end of the lateral line (counted). Scales above lateral line 13(12–14). Scales below lateral line (10 (8–11) Dorsal-fin rays 7(7) Anal-fin rays 6(6) Pectoral-fin rays 11 (10–12) Pelvic 6 (6–7) Caudal-fin rays 19(19), 10 upper rays, 9 lower rays Diagnosis. Separated from all other Speckled Dace by the absence of supraorbital bones (vs. present, Smith et al. 2017). A cryptic species otherwise not distinguishable from other species/subspecies of the Speckled Dace complex except through genomics and geography. Slightly higher lateral line scale counts (mean 74, mode 76, range 64–82) than Sacramento Speckled Dace (mean 70, mode 72, range 54–82) and Lahontan Speckled Dace (mean 68, mode 72, range 60–77) (Table 2). Description. A cryptic species, as shown by genomic analyses; endemic to streams of Los Angeles, San Gabriel, and Santa Ana watersheds in southern California. Recognizable by typical stocky morphology of all Rhinichthys species; small size (adults 6–11 cm SL), wide caudal peduncle (about 1/3 the body depth), blunt-pointed snout, and subterminal mouth. Meristics overlap all other Speckled Dace taxa (Table 2). Body color variable, but adult fish usually pale yellow/white to dusky olive, often with small black spots. Head stripe and lateral spots/blotches pale or absent. Breeding adults of both sexes with orange to red fins; males also with red snouts and lips (Fig. 5). Lateral line usually complete, although last 1–10 scales may lack pores. High overlap in meristic counts with other California Speckled Dace populations (Table 2, Cornelius 1969). Morphometrics. Cornelius (1969) compared the body and fin morphology of 491 dace using 23 truss measurements (divided by standard length or head length, with all values converted to natural logarithms). Only small differences were found between Santa Ana Speckled Dace and dace taxa from other areas. His analysis showed that Santa Ana Speckled Dace had slightly longer fins, more anterior dorsal fins, somewhat longer and narrower heads, smaller eyes, and shorter upper jaws than dace from Sacramento, Lahontan, and Colorado river basins. All differences were not statistically different. However, morphometric differences were sufficient to distinguish lineages when combined meristic measurements and measures of jaw structure (below). Thus, Santa Ana Speckled Dace could be distinguished statistically from nearby Sacramento Basin populations and from Speckled Dace from Lake Tahoe (Lahontan Basin), although overlapping morphometrics made it difficult to assign individuals to any one taxon. Cornelius (1969:26) found that mouth structure provided the strongest separation of the taxa; he used a unique measure of jaw angle: “The lateral angle of the upper jaw formed by the premaxillary and maxillary as measured from photographs of the left side of the head.” Presumably a larger jaw angle is a measure of increased gape and size of prey. Cornelius (1969) found that Santa Ana Speckled Dace had a mean jaw angle of 145 degrees (n = 34, range 128–148) while the Sacramento dace had a mean jaw angle of 160 degrees (n=44, range 146–172). He also found that a frenum is fully developed in 54% (n=277) of the Santa Ana Speckled Dace but in only 2% (n =180) of Sacramento dace. Most (8 of 11 fish examined) Colorado Basin fish had a well-developed frenum. It was absent from (n=41) in Lahontan (Lake Tahoe) dace. Maxillary barbels were present on one or both sides of the mouth in all four dace populations compared, but they were variable in number and size so not useful for distinguishing Santa Ana Speckled Dace. Overall, Cornelius (1969) found that jaw angle, combined with other less quantitative features of the jaw, supported his finding that the Santa Ana Speckled Dace was distinct from dace from Lahontan and Sacramento basins. In combination with other features, jaw structure indicated that Santa Ana Speckled Dace had a common ancestry with fishes from the Colorado River watershed (Virgin River) (Cornelius 1969). This conclusion is supported by genetic and genomic studies (Smith et al. 2002, 2017, Su et al. in 2022). Genomics/Genetics. Genetic studies (mostly using mtDNA) generally indicate that the Santa Ana Speckled Dace is a sister lineage to Speckled Dace populations in the Colorado River Basin as result of a split in lineages approximately 3.6 million years ago (mya) (Oakey et al. 2004, Smith and Dowling 2008). Smith and Dowling (2008), using mtDNA well as morphometrics and meristics, indicate that populations of Speckled Dace in Los Angeles region streams have been isolated long enough (through the Pleistocene) to have diverged from one another and developed some small differences in morphological characters, supporting the findings of Cornelius (1969). The mtDNA analysis by Smith et al. (2017:77) showed “extreme molecular divergence” (about 7%) from other Speckled Dace populations, the highest they encountered. Oakey et al. (2004) also determined that Speckled Dace from the Santa Ana and San Gabriel rivers formed a distinct, monophyletic lineage. Su et al. (2022), using genomic data, confirmed that the Santa Ana Speckled Dace is a sister lineage to dace in the Colorado River Basin. In short, Santa Ana Speckled Dace are quite different genetically from Speckled Dace of other lineages in California and elsewhere (Smith et al. 2017, Su et al. 2022), supporting their designation as a full species (Figure 2). Distribution. Santa Ana Speckled Dace are endemic to three streams, and their tributaries, of the Los Angeles Region in southern California: the Los Angeles, San Gabriel, and Santa Ana rivers. In addition, the San Jacinto River is often treated as a separate river system. However, it historically flowed into the Santa Ana River, a connection severed by dams except in exceptionally wet years. The three river’s headwaters are in the Transverse Mountain Ranges, specifically the San Bernadino, San Gabriel, and Santa Ana mountains. The rivers flowed into an alluvial plain, now covered with Orange County development. During wet years, before the present era, this region would have flooded, connecting the Speckled Dace populations of the rivers. Today the scattered populations of Speckled Dace are largely confined to streams in a band of habitat on public lands that is below the high gradient upper regions of the three watersheds and above the developed alluvial plain. Santa Ana Speckled Dace are absent from many parts of their historic range (Moyle et al. 2015, O’Brien et al. 2011). Each population fragment is increasingly isolated from other populations as the result of dams and other modern infrastructure. Note: Biogeography. There is debate over exactly how the ancestors of Santa Ana Speckled Dace managed to move from the ancestral Colorado River basin to the Los Angeles region (Minckley et al. 1986; Spencer et al. 2008, Smith et al. 2017). They had to survive in a region with a dynamic geology to do so. To colonize coastal California, the dace would have had to travel several hundred kilometers overland (in the modern landscape) over an extended period of time. Speckled Dace in general are freshwater dispersers (Moyle and Cech 2004) so cannot move though marine environments to colonize new areas (Moyle 2002). The dispersal route also has to accommodate two sympatric freshwater dispersers, Santa Ana Sucker (Pantosteus santaanae) and Arroyo Chub (Gila orcutti), both which share ancestry with fishes in the Colorado Basin (Unmack et al. 2014). Smith et al. (2017) argue the fish were able to gradually make their way to California via successive stream captures in a geologically active landscape. This allowed fish to move between headwaters, eventually reaching streams that flowed into the Pacific Ocean from the Coast Range (Santa Monica Mountains). The exact route has been largely erased by the active geology. In contrast, Spencer et al. (2008) present evidence that a likely route was through a combination of two dispersal methods: movement of a plate carrying the fishes followed by colonization via headwater capture. It is likely that dace and other fishes were ultimately moved into the vicinity of the coastal drainages through a combination of the methods proposed by Smith et al. (2017) in concert with the periodic shifting of the main channel of the Lower Colorado River. This channel migration was a dynamic process driven by sediment deposition in the Colorado River channels. Fish presumably followed the river into new channels, including those adjacent to the proto-Santa Ana River and other coastal drainages. The channel migrations eventually blocked the Colorado River, diverting its flow into a large Pleistocene/Holocene lake basin (Lake Cahuilla) subsequent to the formation of the Salton Trough (ca. 1.3 mya). This lake, a predecessor of the Salton Sea, was colonized by lower Colorado River fishes. Bones of the larger fish species have been found in middens of fishing villages along Lake Cahuilla (Wilke 1980). Today, the Lower Colorado River and the Salton Trough are separated from the Santa Ana Basin by the 8–km-high San Jacinto and San Gorgonio Ranges. These ranges are parts of the Peninsula Range and were likely formed by complex plate movements associated with the development a bend in the San Andreas Fault System (ca. 1.1–1.5 mya) (Spencer et al. 2008, Fattaruso et al. 2016). The geologic movements transformed low-relief upland habitat into the dramatic landscape of today. The uplift of the ranges is associated with their westward tilting, resulting in the progressive capture and redirection of eastward-flowing tributaries at a rate of 20–44 km /my (Dorsey and Roering 2006), presumably with the fish. The rapid vertical motions driving these events were dramatic – sometimes exceeding 10m at one time (McClay and Bonora 2001). The suddenness of this redirection of western Colorado River tributaries into coastal-draining basins likely allowed the transfer of freshwater-dispersing fishes into the basin. These fishes apparently became completely isolated no later than ~1.1 mya (Fattaruso et al. 2016, Smith et al. 2017) when uplift of the mountains and downfaulting of the trough created an insurmountable barrier to further dispersal. The uplift the in the Peninsula Ranges (about 1.1–1.5 mya, Fattaruso et al. 2016) created a formidable barrier to freshwater fish dispersal and can be treated as the last point in time that dace could have colonized coastal waters. It is possible that dace were isolated slightly before the San Andreas fault began to shift, as suggested by previous studies (i.e. Spencer et al. 2008), but it is highly unlikely that isolation occurred subsequent to onset of the formation of the Salton Sink (ca. 1.1–1.3 mya) Note: Taxonomy. This study confirms that Santa Ana Speckled Dace is a full species, using the following lines of evidence. Taxonomy: While lacking formal description until now, the Santa Ana Speckled Dace has long been recognized as distinct (Swift et al 1993, Moyle 1976, 2002, Oakey et al. 2004). Definitive morphometric and meristic characters are largely absent. However, when combined, they can be used to distinguish Santa Ana Speckled Dace statistically from populations of other Speckled Dace species/subspecies, as well as from Colorado River Speckled Dace, indicating their long isolation (Cornelius 1969). The apparently unique feature of complete absence of supraorbital bones further supports the species status of this dace. Geology: Improved understanding of the dynamic geology of southern California and of the shifting channels of the Colorado River have resulted in more supportable hypotheses as to how Speckled Dace and other endemic fishes were able to colonize coastal southern California streams. Zoogeography: The historic distribution of the species in streams of the Los Angeles region shows long separation from their ancestral population and long isolation without access to other populations that could lead to hybridization. Although Smith et al. (2017) regard hybridization as the reason R. osculus is just one species throughout its enormous range, including the Los Angeles region, hybridization has clearly not been an issue with the Santa Ana Speckled Dace, which has had no opportunities for over 1.3 million years. Genomics/genetics. Su et al. (2022) show that the Santa Ana Speckled Dace is the most distinct genetically of all other dace populations in California, indicating a long history without gene flow from other dace populations. Etymology. The species epithet honors the Gabrielino-Tongva people, the indigenous inhabitants of the region at the time of the Spanish invasion and who still have a significant presence (https://gabrielinotribe.org/). The original range of the Santa Ana Speckled Dace was in the streams that flowed through the band’s homeland. The tribe is also known as San Gabriel Band of Mission Indians. The common name, Santa Ana Speckled Dace, is widely used for this fish (e.g. Moyle 2002); it shares its name with the Santa Ana Sucker which also occurs in the Santa Ana River and adjacent watersheds. Note: nomenclature. The scientific name has long been a source of confusion. Jordan and Evermann (1896:312) list Agosia nubila carringtoni as having a wide distribution in the Great Basin but state “To this form we also refer provisionally specimens from Lake Tahoe and elsewhere in the Lahontan basin but also those from various coastwise localities in central and southern California, where it is abundant in clear streams and springs as far south as San Luis Obispo.” Agosia n. carringtoni consequently was applied by early workers (e.g., Snyder 1918) to most dace populations in central and southern California including in the first paper mentioning Speckled Dace in the Los Angeles region (Culver and Hubbs 1917). Cornelius (1969:18) summarized the history of the use of carringtoni and quoted a personal communication from Carl Hubbs that there was “no sound basis for continued use of carringtoni in reference to Santa Ana Speckled Dace.” Despite this comment, Cornelius (1969) concluded that R. osculus carringtoni was the only available scientific name. Nevertheless, carringtoni reverted, with no real discussion, to being the name for Speckled Dace found in the Bonneville basin of Utah and Idaho (Smith et al. 2017) from whence it was originally described (Cope 1872). The Santa Ana Speckled Dace has subsequently been referred to as just R. osculus (e.g. Swift et al. 1993, Moyle 1976, 2002) or R. osculus subsp. (e.g., Moyle and Davis 2000, Moyle et al. 2015, Smith et al. 2017) but has resisted formal description until now. Conservation Status. In California, Santa Ana Speckled Dace historically inhabited streams in the upland areas of the San Jacinto, Santa Ana, San Gabriel and Los Angeles rivers which were interconnected by lowelevation floodplains (Swift et al. 1993, Moyle 2002, Moyle et al. 2015) (Figure 5). They have since disappeared from much (75%) of their range, including the middle reaches and tributaries of the Santa Ana River, most of the Los Angeles River and San Jacinto River basins (Moyle et al. 2015, Center for Biological Diversity 2020). Their current distribution is restricted to headwater streams of the San Jacinto, Santa Ana, and San Gabriel rivers and Big Tujunga Creek (Los Angeles River drainage). In 2005, Speckled Dace were reintroduced into the North Fork of Lytle Creek; a similar introduction was made into the Middle Fork in 2007 (Moyle et al. 2015). Both were apparently successful.Attempts to establish populations of Santa Ana Speckled Dace outside their native range were made through introductions into the Santa Clara and Cuyama rivers. Success of these populations is uncertain. An additional introduction into River Springs, Mono County, failed The third edition of the Fish Species of Special Concern Report of the California Department of Fish and Wildlife (Moyle et al. 2015) rates the Santa Ana Speckled Dace as one of the most endangered fish in California. The report notes that the Santa Ana Speckled Dace is a U.S. Forest Service Sensitive Species in the three national forests in which it is found (Angeles, San Bernardino, Cleveland NFs). However, it is likely extirpated from Cleveland National Forest (Santiago Creek). The California Natural Diversity Database ranks Speckled Dace as secure at the global scale (G5) but the Santa Ana subspecies as Imperiled (T1S1; www.natureserve.org). It is listed as Threatened by the American Fisheries Society (Jelks et al. 2008). The habitat of this species continues to shrink due to urbanization of the Los Angeles region, resulting in fragmentation and reduction of populations. Intense human use of their streams on public lands for recreation and water supply also contributes to their decline. Extinction seems likely in the next few decades without special protection for all remaining populations (Swift et al. 1993, Moyle 2002, Moyle et al. 2015). Listing Santa Ana Speckled Dace as threatened or endangered under state and federal Endangered Species Acts would be a major step towards preventing extinction. It appears that lack of a formal species description has been a barrier to listing, even though recent genetic and phylogenetic studies show it to be distinct from all other Speckled Dace (Oakey et al. 2004; Smith and Dowling 2008, Mussmann 2018, Su et al. in 2022). It could therefore be listed as a Distinct Population Segment, even without formal description. Our paper resolves whatever taxonomic uncertainty previously confounded action by regulatory authorities., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 509-517, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Jordan, D. S. & Evermann, B. W. (1896) The Fishes of North and Middle America. Bulletin, US National Museum, 47, Part 1, 1 - 1240.","Culver, G. B. & Hubbs, C. L. (1917) The fishes of the Santa Ana system of streams in southern California, Lorquinia, 1 (11), 82 - 83.","Shapovalov, L. & Dill, W. A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382 - 391","Shapovalov, L., Dill, W. A. & Cordone, A. J (1959) A revised check list of the freshwater and anadromous fishes of California. 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(Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614.","Spencer, J. E., Smith, G. R. & Dowling, T. E. (2008) Middle to late Cenozoic geology, hydrography, and fish evolution in the American Southwest. In: Reheis, M. C., Hershler, R. & Miller, D. (Eds.), Late Cenozoic Drainage History of the Southwestern Great Basin and Lower Colorado River Region: Geologic and Biotic Perspectives. Geological Society of America Special Paper, 439, pp. 279 - 299. https: // doi. org / 10.1130 / 2008.2439 (12)","Moyle, P. B. & Cech, J. J. (2004) Fishes: An Introduction to Ichthyology. 5 th Edition. Prentice-Hall, Upper Saddle River, New Jersey, xvi + 726 pp.","Unmack, P. J., Dowling, T. E., Laitinen, N. J., Secor, C. L., Mayden, R. L., Shiozawa, D. K. & Smith, G. R. (2014) Influence of introgression and geological processes on phylogenetic relationships of western North American mountain suckers (Pantosteus). PLoS One, 9 (3), e 90061. 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Rhinichthys nevadensis robustus (Rutter 1903), new combination, Lahontan Speckled Dace Tables 2, 1g; Figs. 1, 6 Agosia nubila carringtoni Jordan and Evermann 1896:311 Agosia robusta Rutter 1903:148; Rutter 1908:139; Snyder 1917:202 Apocope robusta Snyder 1918:33 Apocope robusta Evermann and Clark 1931:55 Rhinichthys osculus robustus Hubbs and Miller 1948 (a, b) and most subsequent publications. Rhinichthys osculus robustus Hubbs et al. 1974:12 See R. nevadensis for additional synonyms Holotype: USNM 50589, collected from Prosser Creek, California, Rutter and Atkinson (Rutter 1903). Diagnosis. The Lahontan Speckled Dace fits the image of what Speckled Dace are expected to look like (Fig. 1). It is a small (adults usually R. nevadensis by genomics and by its being endemic to the Lahontan Basin in Nevada, Oregon, and California. Description. The following is the original description from Rutter (1903:148) for Lahontan Speckled Dace. “Body heavy, highest above insertion of pectorals; the ventral outline curved almost as much as the dorsal. Head 3.8 to 4 in body; snout blunt, but little overlapping the premaxillary and never extending beyond it; mouth oblique, barbels usually absent, present on 10 to 50 per cent of specimens from any one locality. Fins small; D. 8; A. 7; pectoral about equal to head behind nostril variable; caudal moderately forked, middle rays two-thirds length of longest; rudimentary caudal rays forming prominent keels along upper and lower edges of tail; margin of anal slightly rounded, the anterior rays not all produced, not extending beyond posterior rays when fin is depressed. Lateral line nearly always incomplete, but with scattered pores frequently extending to base of caudal; scales 56 to 77, varying about 12 in any one locality. Usually two dusky lateral stripes, the upper extending from snout to caudal, the lower branching off from the upper behind the head and ending along base of anal; cheek abruptly silvery below lateral stripe; tinged with orange about lower jaw, upper end of gill-opening, and at base of lower fins (p. 148). Note that the coloration described above is based on breeding colors. Hubbs et al. (1974) provide descriptions of the variety of pigmentation patterns present in scattered Speckled Dace populations in the Great Basin and elsewhere. Distribution. Lahontan Speckled Dace are widely distributed in desert basins in northeastern California and northern Nevada (Fig. 3), as described by Hubbs et al. (1974). In California, Rutter (1903) collected specimens from Spring Creek, Willow Creek, Susan River, Little Truckee River, and Prosser Creek, all streams on the western edge of the Lahontan basin. It is also common in Eagle Lake and its inflowing streams (Lassen County) and in the watersheds of the Susan, Truckee, Walker and Carson rivers.The Truckee watershed includes Lake Tahoe and Prosser Creek (tributary to the little Truckee River), the type locality. More broadly, the range of this subspecies includes the drainage of pluvial Lake Lahontan, which covered much of northern Nevada and with fringes in California and Oregon (Hubbs et al. 1974). This region is mostly Great Basin Desert today and includes Pyramid Lake, Nevada, the largest remnant of Lake Lahontan, and the Humboldt River in Nevada. Wherever there is permanent water in the Lahontan Basin, Speckled Dace are likely to be encountered, including isolated populations in spring systems (Hubbs and Miller 1948a, Hubbs et al. 1974, Deacon and Williams 1984). They often co-occur with other Lahontan endemic fishes, such as Tahoe Sucker (Catostomus tahoensis), Lahontan Redside (Richardsonius egregius), Lahontan Tui Chubs (Siphatales bicolor subspp.) and Lahontan Cutthroat Trout (Oncorhynchus clarki lewisi) (Moyle 2002). In recent years, Lahontan Speckled Dace have managed to colonize the headwaters of the North Fork Mokelumne River (Central Valley watershed) as the result of a water project. Lahontan Redsides were also transferred by this same route (Garcia and Associates 2000). Genetics/Genomics. Billman et al. (2010) analyzed mtDNA of dace from the Great Basin; they found support for three distinct lineages (clades): Lahontan, northern Bonneville (Snake River), and southern Bonneville. Smith et al. (2017), using mtDNA, meristics, morphometrics, and fossils, placed all Speckled Dace into three clades, Lahontan, Pacific Northwest (Columbia) and Colorado. Su et al. (2022), using genomics, provide strong support for Lahontan Speckled Dace as a distinct lineage that is a subspecies of Desert Speckled Dace, as are Amargosa Speckled Dace and Long Valley Speckled Dace. Notes. Rutter (1908) found considerable overlap in morphometrics between Lahontan Speckled Dace and dace from the Sacramento drainage, so treated them together as one species. This move was generally not accepted, presumably because the fish faunas of the two basins are otherwise very different. Comparisons of meristics and other features of Lahontan Speckled Dace with those of other dace species and subspecies (including Amargosa and Long Valley Speckled Dace) reflect that there are no external features to easily distinguish the dace subspecies (Tables 2, 3, Figure 1). Our genomic analysis nevertheless shows that Lahontan Speckled Dace is a valid subspecies that is found throughout the Lahontan basin, wherever there is permanent water. The following lines of evidence support this conclusion. Taxonomy. The Lahontan Speckled Dace was described by Rutter (1903) and recognized as a full species until ichthyologist Carl Hubbs and others started calling it a subspecies, Rhinichthys osculus robustus. This designation was justified by the lack of features easily separating it from other Speckled Dace taxa. Geography/Geology. The distribution of Lahontan Speckled Dace coincides with the northern part of the region called the Great Basin Desert (Deacon and Williams 1984; Moyle 2002). Genomics/genetics. Three genetic/genomic studies have shown that Lahontan Speckled Dace belong to a distinct evolutionary lineage as do the two other subspecies of Desert Speckled Dace (Oakey et al. 2004; Billman et al, 2010; Su et al., 2022). The long isolation of this subspecies in the region is reflected in the lack of genomic evidence for recent hybridization with other dace species/subspecies. Etymology. The Lahontan Speckled Dace was originally described by Rutter (1903) as Agosia robusta without explanation of the species name, although he described the body as heavy, perhaps reflecting the roundness of the body in cross-section, the blunt snout, and thick caudal peduncle. Gilbert (1893) also described the Amargosa Speckled Dace as being robust. Conservation Status. The Lahontan Speckled Dace is widespread and abundant in many places in the Great Basin Desert and in streams and lakes of the Eastern Sierra Nevada. Because streams are increasingly dammed and diverted for human use, dace populations are increasingly fragmented. Speckled Dace are rare in or absent from most reservoirs. The safe status of Lahontan Speckled Dace should therefore not be taken for granted, especially in the face of climate change., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 519-520, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Rutter, C. (1903) Notes on fishes from streams and lakes of northeastern California not tributary to the Sacramento basin. United States Fish Commission Bulletin, 22, 145 - 148. [1902]","Jordan, D. S. & Evermann, B. W. (1896) The Fishes of North and Middle America. Bulletin, US National Museum, 47, Part 1, 1 - 1240.","Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin, with a study of their distribution and variation. United States Bureau of Fisheries Bulletin, 27, 105 - 152. [1907]","Snyder, J. O. (1917) An account of some fishes from Owens River, California. Proceedings, United States National Museum, 54, 201 - 205. https: // doi. org / 10.5479 / si. 00963801.2233.201","Snyder, J. O. (1918) The fishes of the Lahontan system of Nevada and northeastern California. United States Bureau of Fisheries Bulletin, 35, 33 - 86.","Evermann, B. W. & Clark, H. W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1 - 67.","Hubbs, C. L., Miller, R. R. & Hubbs, L. C. (1974) Hydrographic history and relict fishes of the North-central Great Basin. San Francisco: California Academy of Sciences Memoir, 7, 1 - 259.","Hubbs, C. L. & Miller, R. R. (1948 a) The Great Basin, with emphasis on glacial and postglacial time. The zoological evidence: correlation between fish distribution and hydrographic history in the desert basins of western United States. University of Utah Bulletin, 38 (20), 17 - 166.","Deacon, J. E. & Williams, J. E. (1984) Annotated list of the fishes of Nevada. Proceedings of the Biological Society of Washington, 97 (1), 113 - 118.","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Garcia & Associates (2000) The Distribution and Abundance of the Benthic Macroinvertebrate Fauna and Fish Populations in Tributaries Leading into and including the North Fork Mokelumne and Mainstem Mokelumne Rivers. Final Report Volume 3. Prepared for Pacific Gas and Electric Company by Garcia and Associates, San Anselmo, California. [unknown pagination]","Billman, E. J., Lee, J. B., Young, D. O., McKell, M. D., Evans, R. P. & Shiozawa, D. K. (2010) Phylogenetic divergence in a desert fish: differentiation of Speckled Dace within the Bonneville, Lahontan, and upper Snake River basins. Western North American Naturalist, 70 (1), 39 - 48. https: // doi. org / 10.3398 / 064.070.0105","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Oakey, D. D., Douglas, M. E. & Douglas, M. R. (2004) Small fish in a large landscape: diversification of Rhinichthys osculus (Cyprinidae) in western North America, Copeia, 2004 (2), 207 - 221. https: // doi. org / 10.1643 / CG- 02 - 264 R 1","Gilbert, C. H. (1893) Report on the fishes of the Death Valley expedition collected in southern California and Nevada in 1891 with descriptions of new species. U. S. Department of Agriculture, North American Fauna, 7, 229 - 234. https: // doi. org / 10.3996 / nafa. 7.0003"]}

Rhinichthys nevadensis caldera, new subspecies, Long Valley Speckled Dace Table 3, Fig. 4. Synonymy: Rhinichthys osculus subsp. Sada et al. 1995:356; Moyle 2002:161. Otherwise see synonyms for Desert Speckled Dace, Rhinichthys nevadensis. Holotype: WFB 5000 (Fig. 4). Standard length 60 mm, fork length 70 mm; White Mountain Research Center, Northeast Pond 37.360618°N 118.329370°W. Inyo County, California, July 29, 2021. Nicolas Buckmaster, Rosa Cox, California Department of Fish and Wildlife Paratypes: WFB 5001-5009 (n=9). Same as holotype. Meristic s: holotype (paratypes) Lateral line scales: 64 (59–80). Lateral line incomplete. Scales above lateral line: 10 (9–12), scales difficult to discern. Scales below lateral line: 9 (6–10), scales hard to discern Dorsal-fin rays: 8(7–8), counts include single unbranched ray. Anal-fin rays: 7 (6–7), counts include single unbranched ray. Pectoral-fin rays 13 (11–12), counts include anterior and posterior single unbranched rays. Pelvic 7 (6–8), counts include single unbranched ray. Caudal-fin rays 19 (19). Diagnosis. The Long Valley Speckled Dace is a cryptic taxon, very similar in its meristics to the Amargosa Speckled Dace (Table 3). They are distinguished from other Speckled Dace by their being endemic to the Long Valley region and having a distinct evolutionary lineage, as revealed by genomics (Su et al. 2022). Adults are small (rarely more than 75 mm SL) and recognizable as Speckled Dace by their thick caudal peduncle, sub-cylindrical body, small fins, small eyes (relative to head), and blunt, pointed snout. In life, they are a bronzy color with light speckling (Fig. 6). Striping is largely absent. Maxillary barbels and a frenum are usually absent. They can be distinguished statistically from similar Amargosa Speckled Dace by slightly higher average numbers of pectoral and pelvic, higher lateral line scale count, lower lateral line pore scale count, and absence of maxillary barbels (Sada et al. 1989, 1995). The following mean counts (standard error) are from Long Valley Speckled Dace collected in Whitmore Hot Springs and at an unnamed spring at Little Alkali Lake (Sada 1989, Table 3): lateral line scales 61.7 (1.4); lateral line scales with pores 19.0 (5.0); dorsal-fin rays 8.0 (0.0); anal-fin rays 7.0 (0.0); pectoral-fin rays 13.0 (0.4); pelvic 7.4 (0.2). Description. Long Valley Speckled Dace are small, active fish with the typical Speckled Dace morphology, as described for the species. Genetics/Genomics. Sada et al. (1989, 1995), using isozymes and morphometrics, were the first to recognize Long Valley Speckled Dace as different from other dace populations in the region. Oakey et al. (2004), Mussmann et al (2020) and Su et al (2022) all found genetic differences that separated Long Valley Speckled Dace from Amargosa and Lahontan Speckled Dace. Distribution. The historic range of this dace was in the outlets of hot springs and associated marshes in the remains of the Long Valley volcanic caldera, just east of Mammoth Lakes, Mono County, as well as in Hot Creek. It quite likely had its origins when Speckled Dace colonized the upper Owens Valley region from the Mono Lake Basin via Adobe Valley, during a late Pleistocene pluvial period. During this time, Mono Lake levels were high enough so that it periodically spilled into Adobe Valley, which drained into the Owens River, from which fish presumably moved upstream into Long Valley. Subsequent down-faulting of the Owens River and formation of steep waterfalls in the Owens River gorge likely isolated Long Valley from the Owens Valley around 100,000 years ago (Hildreth and Fierstein 2016). Note. Long Valley Speckled Dace were considered to be one of the scattered populations of Owens Valley Speckled Dace until Sada et al. (1993, 1995) showed it was genetically and morphometrically distinct and that it was closely related to dace populations in Ash Meadows, Amargosa River, and elsewhere in Owens Valley and Death Valley. Moyle (2002) and Moyle et al. (2015) recognized it as an undescribed subspecies. Our analysis shows that the Long Valley Speckled Dace is a distinct lineage confined to a small part of the Owens Valley region. It merits subspecies designation based on the following lines of evidence. Taxonomy. Multiple analyses indicate that Long Valley dace is a distinct lineage (subspecies) within Desert Speckled Dace, R. nevadensis (Oakey et al 2004, Mussmann et al. 2020, Sada et al. 1995, Moyle 2002, Moyle et al. 2015 and Su et al., 2022). Geography/geology. During pluvial periods of the Pleistocene, the Death Valley region was a series of interconnected large lakes, with abundant fishes (Hubbs and Miller 1948). When the climate changed and heavy precipitation stopped falling, the lakes and rivers dried up or became small remnants of what they once were. This process resulted in numerous isolated Speckled Dace populations with little, or at least very infrequent, opportunity for genetic exchange. See Desert Speckled Dace account for more details. Long Valley is part of the Death Valley regional endemism hotspot with numerous endemic plants and animals, including fishes (Sada et al. 1995). Long Valley Speckled Dace can be regarded as another organism endemic to the isolated Owens Valley region. Genetics/genomics. Sada et al. (1993, 1995) were first to recognize that Long Valley Speckled Dace were genetically distinct from other dace, a finding confirmed by other studies (Mussmann et al. 2020, Su et al. 2022). See Amargosa Speckled Dace (R. nevadensis nevadensis) description for more details. Etymology. The Long Valley Speckled Dace was historically known only from small streams flowing into the Long Valley Caldera, the remnants of a gigantic volcano that last erupted 0.7 million years ago (Hildreth and Fierstein 2016). They colonized the remnants of the caldera during the late Pleistocene, hence caldera. Conservation Status. Long Valley Speckled Dace have been extirpated from all but one of their historic collection sites, including Hot Creek. The only population left in its native range (as of 2021) is in Whitmore Marsh and its inlet stream in Long Valley (Moyle et al. 2015). Unfortunately, this marsh is now maintained by the outflows of a hot spring system that has been developed as a public swimming pool by the Town of Mammoth Lakes. Discharge is approximately 2 cfs and is lightly chlorinated. The outlet stream feeds an alkaline marsh of roughly 1 acre. In 1989, dace occupied 250 meters of stream and two large shallow ponds less than a half meter deep (Moyle et al 2015). Surveys in 2002 and 2009 by CDFW found this population to be relatively stable (S. Parmenter, CDFW, pers. comm. 2009). In 2018, 2019, and 2020, they were not found in these habitats (S. Parmenter, CDFW, in Center for Biological Diversity, 2020). However, in 2021 and 2022, trapping produced a few fish (R. Black, CDFW, pers. comm, 2021). The only other population is in a single artificial pond into which they were introduced at the White Mountain Research Center, outside their native range; the status of this dace population is checked by CDFW on a routine basis (R. Black, CDFW, pers. comm.). The multiple causes of decline are discussed in Moyle et al. (2015) and Center for Biological Diversity (2020). The CBD (2020) has petitioned to have this dace listed as endangered under the federal ESA or else have all Speckled Dace in the Death Valley region included under the 1984 listing of Ash Meadows Speckled Dace as endangered. It is listed as a Species of Special Concern (critical concern) by CDFW.

Rhinichthys klamathensis klamathensis, new combination, Klamath Speckled Dace, Fig. 6 Synonymy. Same as for Western Speckled Dace, R. klamathensis. Holotype and paratypes are the same as for Western Speckled Dace, R. klamathensis. Diagnosis. Same as Western Speckled Dace, R. klamathensis; the Speckled Dace lineage that is abundant in the streams, lakes, and other aquatic habitats in the Klamath River Basin in Oregon and California, including the Trinity River in California. Distinguished statistically by having somewhat smaller scales (scales in lateral line, 68–78; mean 73, n =49) than the other two subspecies (Sacramento, 54–83, mean 70, n=123; Warner, 68–71, mean 69, n=58). Otherwise, it is not distinguishable from other Speckled Dace except by genomics and distribution. Description. Evermann and Meek (1898) described this cryptic subspecies as Agosia klamathensis. Therefore, the description quoted in the Western Speckled Dace account is also for this subspecies, and it applies to the other two subspecies as well. Distribution. The Klamath Speckled Dace is found throughout the Klamath Basin in Oregon and California, including the Klamath and Trinity rivers and tributaries, as well as in the Rogue River to the north (Wiesenfeld et al. 2017). It is also widespread in the upper Klamath Basin, including Upper Klamath Lake. The Speckled Dace in the Eel River, to the south of the Klamath Basin, is an introduced population of R. k. klamathensis (Kinziger et al. 2011). Geology/zoogeography. The geologic history of the Klamath region is complex. The Snake River, now a tributary to the Columbia River, was once a major river that originated in the Idaho region and flowed to the ocean via the ancestral Klamath River during the Pliocene (Minckley et al. 1986, Smith et al 2017). When the Snake broke through to the Klamath, it carried with it the lake fauna of the Great Basin, which gave rise to the presentday endemic, freshwater-dispersing fish fauna (12 species) of the upper Klamath basin (Moyle 2002), presumably including Speckled Dace. Pfrender et al. (2004:498), using a molecular clock based on mtDNA from Speckled Dace from the major river basins of Oregon, speculated that “…the levels of sequence divergence in R. osculus among these major basins are more consistent with a Pliocene or Miocene sundering of gene flow between major basins. Miocene isolation of these river systems is substantially earlier than has previously been suggested.” They thought such ancient divergence could explain the genetic diversity found in dace within the Klamath basin, a diversity that was also documented by Wiesenfeld et al. (2017). See also the geology discussion under Western Speckled Dace. Geographically, Klamath Speckled Dace co-occur with other fish species endemic to the Klamath watershed. However, the endemic fishes in the upper watershed (above Klamath Falls) are different from those in the lower river and it appears that dace from the two regions diverge as well, a situation noted also for Marbled Sculpin, Cottus klamathensis (Daniels and Moyle 1984). Genetics/genomics. The genomic study of Su et al. (2022) unveiled the three lineages within R. klamathensis, as discussed. Within the Klamath drainage, Wiesenfeld et al. (2017:8), using analyses of microsatellites and mtDNA, found that The Klamath–Trinity Basin Speckled Dace were resolved as nearly monophyletic [bootstrap (BS) 77], and exhibited a sister group relationship with nearby basins in California and Oregon (Sacramento, Pit River, and Goose Lake). The Klamath River and Trinity River populations, however, were found to be sufficiently distinct to suggest the two rivers were once isolated from one another, with their connection being fairly recent. The presence of apparent hybrids in the region where the two rivers meet today also supports this conclusion (Wiesenfeld et al. 2017). A genetically distinct population is also found in Jenny Creek, a tributary to the Klamath River that is largely in Oregon (Wiesenfeld et al. 2017). Pfrender et al (2004) used mtDNA to compare dace from the upper Klamath and Rogue rivers with those from two tributaries to the Columbia River in Oregon. They found species-level differences between dace in the two great watersheds, with estimated divergence times of 3.9 to 9.6 million years ago. Within the upper Klamath basin, Pferender et al. (2004) found considerable genetic structure but of more recent origin (et al. (2017). Note. The Klamath Speckled Dace was originally described as a species that differed from other Speckled Dace by having finer scales (Evermann and Meek 1898). Subsequently, it retained its identity as a subspecies, R. osculus klamathensis. This is not surprising given that the Klamath Basin has long been recognized as an isolated basin which supports high endemism in its fishes (Moyle 2002). The Klamath Speckled Dace has retained its identity through the region’s complex geologic history, enduring the rise of mountain ranges, high levels of vulcanism, and invading interior rivers (Minckley et al. 1986). This complex geologic history has kept the Klamath fish fauna isolated and distinct. In the upper Klamath basin, Speckled Dace live with other endemic fishes that had their ancestors in the Great Basin. Even in the lower Klamath River, which is too swift for the lake-adapted fishes of the upper basin, it co-occurs with the endemic Klamath Smallscale Sucker, Catostomus rimiculus (Moyle 2002). Given the genetic structure that Wiesenfeld et al. (2017) found in the basin, it is possible that dace populations from the upper and lower river should be treated as separate subspecies. Su et al. (2022) confirmed results from other genetic methods that the Klamath Speckled Dace is a distinct evolutionary lineage. Etymology. The Klamath Speckled Dace is named for the river system to which it is endemic. The river is named for the native peoples who lived (and still do) in the upper Klamath Basin. Conservation Status. The Klamath Speckled Dace is widespread and abundant in streams and natural lakes in the Klamath Basin, including the Trinity River. The main concern is that some distinct populations, such as in Jenny Creek, may be lost as the waters are dammed and diverted. The genetic diversity of Speckled Dace populations within the river system is just beginning to be appreciated (Pfrender et al. 2004; Wiesenfeld et al. 2017)., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 526-528, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Evermann, B. W. & Meek, S. B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U. S. Fish Commission, 17, 15 - 84. https: // doi. org / 10.5962 / bhl. title. 35601","Rutter, C. (1903) Notes on fishes from streams and lakes of northeastern California not tributary to the Sacramento basin. United States Fish Commission Bulletin, 22, 145 - 148. [1902]","Kinziger, A. P., Nakamoto, R. J., Anderson, E. C. & Harvey, B. C. (2011) Small founding number and low genetic diversity in an introduced species exhibiting limited invasion success (Speckled Dace, Rhinichthys osculus). Ecology and Evolution, 1 (1), 73 - 84. https: // doi. org / 10.1002 / ece 3.8","Minckley, W. L., Hendrickson, D. A. & Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. & Wiley, E. O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614.","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Pfrender, M. E., Hicks, J. & Lynch, M. (2004) Biogeographic patterns and current distribution of molecular-genetic variation among populations of Speckled Dace, Rhinichthys osculus (Girard). Molecular Phylogenetics and Evolution, 30 (3), 490 - 502. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00242 - 2","Daniels, R. A. & Moyle, P. B. (1984) Geographic variation and a taxonomic reappraisal of the Marbled Sculpin, Cottus klamathensis. Copeia, 1984, 949 - 959. https: // doi. org / 10.2307 / 1445339","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388"]}

Rhinichthys klamathensis goyatoka, new subspecies. Warner Speckled Dace Fig. 4C Agosia nubila carringtoni Snyder 1908:98 Agosia robusta Rutter 1908: 139 Apocope carringtoni Evermann and Clark 1931:55 Rhinichthys osculus subsp. Williams et al. 1990:243 Rhinichthys osculus klamathensis Markle 2016:50 Holotype: WFB-122-10-44 (Figure 1). 59 mm SL. Twelve Mile Creek, Lake County, Oregon. Jack E. Williams. November 2, 1988. Paratypes: WFB-122-10-42a—122-10-42j (n = 10 ). Same location and collector Meristics: Holotype (paratypes) Length (mm): standard 59; fork 67; total 71 Lateral line scales: 76 (61–84). Lateral line incomplete in most individuals; counts include 2–3 scales beyond end of lateral line. Scales above lateral line: 13 (11–14) Scales below lateral line: 11 (9–13) Dorsal-fin rays: 8(8), counts include single unbranched ray. Anal-fin rays: 7 (7), counts include single unbranched ray. Pectoral-fin rays 12 (10–13), counts include unbranched rays. Pelvic 7 (7–9), counts include unbranched rays. Caudal-fin rays 19 (19–20). Diagnosis. Genetically distinct Speckled Dace endemic to the isolated Warner Basin in Oregon and California. Relatively small adult size (Description. The description is the same as Sacramento Speckled Dace. Snyder (1908) examined Speckled Dace from throughout the arid basins of southeastern Oregon and could find no distinguishing morphological or meristic features to separate Warner Speckled Dace from dace in other basins, not even lateral line scale numbers (which partially define Klamath Speckled Dace). For lateral line scales, Snyder’s counts were: Warner Basin 68– 71(N=58), Sacramento, 61–74 (N=143), and Klamath 68–78 (n=18). He also noted that in Warner Speckled Dace, maxillary barbels were usually present (45 out of 53 fish examined). For comparison, they were present on 43/48 Klamath basin dace but only 9/68 for Sacramento basin dace. A frenum is usually present. Distribution. In a survey of fishes of the Warner Basin, Williams et al. (1990) found Speckled Dace in Twelvemile Creek, Twentymile Creek, Deep Creek, and upper Honey Creek, plus the isolated Foskett Spring. Only Twelvemile Creek has headwaters (and Speckled Dace) in California. Foskett Spring is on the edge of Coleman Lake, on the southeast corner of the basin; the lake is dry most of the time. The mtDNA study by Smith et al. (2017) indicates Speckled Dace in the Warner Basin share a recent ancestry with those from the Chewaucan River to the north (which flows into Lake Abert, Oregon) and with those in Wall Canyon Creek (Nevada) to the southeast. Further genomic studies may show these populations could be included within Warner Speckled Dace. Geology. The Warner Valley is one of a number of isolated watersheds within the northern Great Basin, which have a long complex history of occasional connectivity to big river systems, such as the Snake and the Klamath (Smith et al. 2002). The onset of Great Basin faulting and uplift of the Warner Range (ca. 3 mya) likely resulted in a permanent separation and isolation of the Warner Basin from the Sacramento and Klamath basins, sometime in the late-Pleistocene (~1.0 mya) (Eggar et al. 2011). This isolation is reflected in other endemic fishes which co-occur with Warner Speckled Dace: Warner Sucker (Catostomus warnerensis), Great Basin Redband Trout (Oncorhynchus mykiss subsp.) and Tui Chub (Siphatales bicolor subsp.) (Williams et al. 1990; Markle 2016). In fact, the Warner Valley is one of a number of isolated watersheds within the northern Great Basin that support endemic aquatic species (Hubbs et al. 1978), so it is likely that Speckled Dace are endemic to other watersheds as well. Genetics/genomics. Ardren et al. (2010), using mtDNA, suggested that Speckled Dace in the Warner Basin are different at the species level from dace in the neighboring Goose Lake Basin (Sacramento drainage). Hoekzema and Sidlauskas (2014) examined mtDNA from dace populations in the Warner Basin and in surrounding basins in Oregon. They found a high level of genetic divergence in dace from the basins, dating to the Pliocene or early Pleistocene, which “…should likely be elevated to species-status once their full geographic extent is discovered, and their morphological diversity described (Hoekzema and Sidlauskas 2014:245). Distinct lineages included fishes from Goose Lake, Silver Lake, Lake Abert, and the Warner basin, including Foskett Spring. The mtDNA study of Smith et al. (2017) and the genomic study of Su et al. (2022) both indicate that Warner Basin Speckled Dace are a distinct lineage, divergent from the Sacramento and Klamath lineages, at the subspecies or species level. Etymology. The name honors the Goyatöka people who, before the invasion of their homeland by Euro- Americans, lived on lands in the Warner Basin where Speckled Dace occupied the streams, lakes, and springs (Dixon 1908, Stewart 1939). The common name indicates the geographic location of the dace populations. The basin is named for William Warner, an army officer who died while mapping the region. Notes. Our study supports the finding that the Warner Speckled Dace is a distinct lineage (subspecies) allied with Klamath and Sacramento Speckled Dace. Ardren et al. (2010) and Su et al. (2022) show that Warner Speckled Dace have evolutionary ties to the daces in the Klamath and Sacramento Basins. These Speckled Dace became a species of interest when dace in Foskett Spring, in an isolated part of the Warner Basin, were listed as a threatened species without any special taxonomic designation. Subsequent genetic analyses revealed Foskett Spring Speckled Dace could at best be considered a Distinct Population Segment of Warner Speckled Dace; these same analyses suggested that Speckled Dace throughout the Warner Basin constituted a distinct taxon (Ardren et al. 2010). Conservation Status. Most of what is known about the Warner Speckled Dace and other fishes of the basin comes from surveys for (a) Warner Sucker, which was listed as threatened in 1985, (b) Foskett Spring Speckled Dace, which was also listed as a threatened species in 1985 (USFWS 2019, https://ecos.fws.gov/ecp/species/651), and (c) Great Basin Redband Trout. The Foskett Spring Speckled Dace was delisted in 2019 as the result of extensive habitat improvement, although it remains a conservation dependent species (USFWS 2019). Most of the permanent water in the basin inhabited by the Warner Sucker is dominated by non-native fishes, but a population of native Tui Chub is also present (Williams et al. 1990). The remaining Warner Speckled Dace populations (and those of Warner Sucker and Tui Chub) probably qualify for listing as Threatened under the federal ESA. Their decline is due to the combined problems of non-native predatory fishes occupying much of the aquatic habitat, inadequate management of grazing and diversions, and climate change/drought reducing flows and increasing temperatures in streams (Williams et al. 1990)., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 531-532, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Snyder, J. O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U. S. Bureau of Fisheries Bulletin, 27, 69 - 102.","Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin, with a study of their distribution and variation. United States Bureau of Fisheries Bulletin, 27, 105 - 152. [1907]","Evermann, B. W. & Clark, H. W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1 - 67.","Williams, J. E., Stern, M. A., Munhall, A. V. &. Anderson, G. A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243 - 248. [http: // www. jstor. org / stable / 41712597]","Markle, D. F. (2016) A Guide to the Freshwater Fishes of Oregon. Oregon State University Press, Corvallis, Oregon, 140 pp.","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Smith, G. R., Dowling, T. E., Gobalet, K. T., Lugaski, T., Shiozawa, D. & Evans, P. (2002) Biogeography and timing rates of evolutionary events among Great Basin fishes. In: Hershler, R., Madsen, D. B. & Curry, D. R. (Eds.), Great Basin Aquatic Systems History, Smithsonian Contributions to the Earth Sciences, 33, pp. 175 - 233.","Ardren, W. R., Baumsteiger, J. & Allen, C. S. (2010) Genetic analysis and uncertain taxonomic status of threatened Foskett Spring Speckled Dace. Conservation Genetics, 11 (4), 1299 - 1315. https: // doi. org / 10.1007 / s 10592 - 009 - 9959 - 0","Hoekzema, K. & Sidlauskas, B. L. (2014) Molecular phylogenetics and microsatellite analysis reveal cryptic species of Speckled Dace (Cyprinidae: Rhinichthys osculus) in Oregon's Great Basin. Molecular Phylogenetics and Evolution, 77, 238 - 250. https: // doi. org / 10.1016 / j. ympev. 2014.04.027","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Dixon, R. B. (1908) Notes on the Achomawi and Atsugewii Indians of northern California. American Anthropologist, 10 (2), 208 - 220. https: // doi. org / 10.1525 / aa. 1908.10.2.02 a 00030","Stewart, O. C. (1939) The Northern Paiute Bands. Anthropological Records, 2 (3), 127 - 149."]}

Rhinichthys nevadensis nevadensis Gilbert 1893 new combination, Amargosa Speckled Dace. Table 3. Synonymy. See Desert Speckled Dace, Rhinichthys nevadensis, account. Holotype. See Desert Speckled Dace, Rhinichthys nevadensis, account. Diagnosis. The Amargosa Speckled Dace is a cryptic taxon diagnosable with genomics and distribution as the Speckled Dace found in the springs, streams, and ditches of Death Valley, Owens Valley, and the Amargosa River. Individuals are typically small (8–11cm SL) and recognizable as Speckled Dace by their thick caudal peduncle, sub-cylindrical body, small fins, small eyes, and blunt pointed snout. The sides of the body are usually covered with black blotches which merge into a black stripe, which is especially dark below the eye. Gilbert’s (1893:230) short diagnosis is: “Differing from other known species [of dace] by the large head, the short deep body, very small eye, and in the reduction of the outer ventral [fin] ray to a mere rudiment.” He also noted the presence of an incomplete lateral line, of maxillary barbels, and of a conspicuous lateral stripe. See Table 3 for meristics. However, these characters, except genomics and distribution, do not separate the three R. nevadensis subspecies from one another. Description. The formal description of Amargosa Speckled Dace (as a species) by Gilbert (1893) is thorough (Figure 6) and is included in this paper as the description of Desert Speckled Dace, R. nevadensis. Scoppettone et al. (2011) measured 1354 dace from the Amargosa River Canyon, which had a mean fork length of 51mm and a maximum FL of 92 mm FL. None of Gilbert’s characters separate the regional Speckled Dace populations from one another, demonstrating the cryptic nature of Speckled Dace taxa. Sada et al. (1995) performed a principal component analysis on 20 morphological measurements (truss analysis) and eight meristic characters on 484 Speckled Dace (386Amargosa Speckled Dace and 98 Lahontan Speckled Dace) from 13 localities (Table 3); they found that “Meristic and proportional mensural characters were all within ranges documented for Speckled Dace…Highly significant differences among all populations for all characters showed the morphology of each population to be unique (Sada et al. 1995:352–353).” Essentially, they found evidence of local adaptation and genetic drift, but no features that could readily distinguish individual Amargosa Speckled Dace from individuals in other regional dace populations, including Lahontan and Long Valley Speckled Dace. Lahontan Speckled Dace, however, did have slightly more lateral line scales than the other two subspecies but overlap in scale counts and incomplete lateral lines in many individuals makes this character unreliable for identification (Table 3). Distribution. Gilbert (1893) reported the distribution of R. nevadensis as warm water springs in Ash Meadows, Indian Creek, and Vegas Creek, Nevada. Vegas Creek has been covered up by Las Vegas and its endemic Speckled Dace, Rhinichthys deaconi, is extinct (Smith et al. 2017). The Amargosa Speckled Dace is endemic to springs, small streams, and rivers in the Death Valley system (which includes the Owens Valley) in California and Nevada (Figure 3). In the Owens Valley, it is assumed they once were found throughout the length of the Owens River (Snyder 1917). For more details of distribution see the Notes section of this account. Genetics/genomics. Early genetic studies generally concluded that Speckled Dace from the Death Valley region were all one lineage but included branches variously designated as Owens, Ash Meadows, Amargosa River, and Long Valley Speckled Daces. Thus, Sada et al. (1993, 1995) used protein electrophoresis to examine relationships among dace from 13 localities in the region and from the once-interconnecting Amargosa River. They found that all populations had some statistically identifiable features and that the regional Speckled Dace populations likely diverged from Lahontan Speckled Dace after vulcanism and drying-up of interconnecting waterways isolated them in the Death/Owens Valley region. Oakey et al. (2004), using mt/DNA found that all populations of Speckled Dace in the region formed a clade. The genomic analysis of Mussmann et al. (2020) produced similar results. They recommended that the populations be treated as Distinct Population Segments within the Amargosa Speckled Dace lineage because they met three criteria: current geographic isolation, genetic differentiation, and local adaptation. According to Mussmann et al. (2020:13) “the identities of the various lineages is intimately tied to the prehistoric lakes and rivers of the region, with diversifications occurring within modern drainages. This pattern clearly reflects a relictual biodiversity with high endemicity, with patterns driven by Plio-Pleistocene tectonics and hydrology (i.e., dispersal of Speckled Dace from Owens Valley to the Amargosa Basin during fluvial events)…”. The analysis of Su et al (2022) agrees with this statement and recognizes that the Desert Speckled Dace includes three subspecieslevel lineages (Lahontan, Amargosa, and Long Valley) with the Lahontan lineage likely being the most immediate ancestral population. Notes: Subspecies. The Amargosa Speckled Dace is here designated as a subspecies of Desert Speckled Dace (Rhinichthys nevadensis), going back to Gilbert’s original description and name, based on the following lines of evidence: Taxonomy. The Amargosa Speckled Dace has been a stable taxon since it was described, although it was mysteriously switched from being a species to a subspecies of Rhinichthys osculus in the 1940s, following the Great Basin studies of Hubbs and Miller (1948) and Miller (1946). Gilbert’s original description only encompassed dace in Ash Meadows, leaving as undescribed other dace populations in the Amargosa River, Oasis Valley, and Owens Valley. The undescribed forms have been variously lumped with Amargosa (Moyle 1976) or Lahontan Speckled Dace (Snyder 1917, 1918) or treated as undescribed subspecies of R. osculus (Miller 1973). Moyle et al. (2015) divide the Death/Owens Valley daces into Long Valley, Owens, and Amargosa populations. Su et al. (2022), however, show that the Owens Valley populations are best treated a part of the Amargosa subspecies, R. nevadensis nevadensis. Zoogeography. During pluvial periods of the Pleistocene, the Death Valley and Lahontan regions supported many interconnected large lakes, with abundant fishes (Hubbs and Miller 1948). When the climate changed and frequent precipitation stopped, the lakes and rivers dried up or became small, disconnected remnants of what they once were. As a result, the Death Valley region became an endemism hot spot with numerous endemic plants and animals in springs and wetlands (Sada et al. 1995). The fish that survived became restricted to relatively small areas where permanent stream flows were created by springs, which is where we find them today. For Speckled Dace, this process resulted in numerous isolated populations with little opportunity for genetic exchange in the present landscape. In Death Valley proper, including Ash Meadows, the Speckled Dace shares spring systems with four species and eight subspecies of pupfish (Cyprinodon spp.). In the Owens Valley, other endemic fishes are Owens Tui Chub (Siphatales bicolor snyderi), the undescribed Toikona Tui Chub (Chen et al. 2007), Owens Sucker (Catostomus fumeiventris) and Owens Pupfish (Cyprinodon radiosus). Genomics. Su et al. (2022) support Amargosa Speckled Dace as a distinct lineage within Desert Speckled Dace. Note: Distinct Population Segments. The Federal Endangered Species Act of 1973 defines species fairly broadly for fishes and other vertebrates. The act recognizes that populations that are distinct and important segments (usually defined by genetics and distribution) of a species total population can be listed as Threatened or Endangered, even if the species is not so listed. To be recognized, Distinct Population Segments (DPSs) do not go through a formal taxonomic description process. Thus, Mussmann et al. (2020) used genomics to determine that the populations of Desert Speckled Dace could be divided into three DPSs: Long Valley, Owens River, and Oasis Valley. “Ash Meadows and Amargosa River populations are included within the Oasis Valley DPS. In addition, they recognized that dace in Amargosa Canyon are of hybrid origin (Amargosa X Ash Meadows). We treat this fish as a population of Amargosa Speckled Dace. Our analysis agrees with Mussmann et al., (2020) that the Long Valley Speckled Dace are sufficiently different to be recognized as a subspecies. However, our analysis also recognizes Ash Meadows and Owens Valley Speckled Dace as two DPS’s within Amargosa Speckled Dace. The Ash Meadows Speckled Dace was first described from springs and their outflows in Ash Meadows. Genetically similar fish occupy similar habitat in the Oasis Valley, Nevada. In some years, the water from both areas ultimately flows into the Amargosa River and down into Amargosa Canyon. Therefore, this DPS includes daces from Ash Meadows, Oasis Valley, and Amargosa Canyon, including Willow Creek (Mussmann et al. 2020). The Owens Valley Speckled Dace is found only in streams, springs and ditches of the Owens Valley, California. It was considered by La Rivers and Trelease (1962) and Moyle (1976) to be a population of Amargosa Speckled Dace (R. osculus nevadensis). Sada (1989) and Sada et al. (1993, 1995) showed it was genetically and morphometrically separable from populations in Ash Meadows, Amargosa River, and Long Valley. Moyle (2002) and Moyle et al. (2015) recognized it as an undescribed subspecies related to Amargosa Speckled Dace. Otherwise, synonymies are as for the species (R. nevadensis) and subspecies (R. n. nevadensis, R. n. caldera). Etymology. This subspecies, as the species Rhinichthys (Apocope) nevadensis, was described by Gilbert (1893:230) from “Ash Meadows, Amargosa Desert, on boundary between California and Nevada” hence the common and scientific names. The springs in Ash Meadows ultimately drain into the Amargosa River, which flows into Death Valley. Amargosa is derived from a Spanish word meaning bitter and refers to the unpalatability of the water. Jordan and Evermann subsequently (1896) placed the species in Agosia. The members of this genus were returned to Rhinichthys in the 1940s and the species simultaneously became a subspecies of R. osculus, both without a formal process. Conservation Status. All populations of R. n. nevadensis are threatened with extinction to a greater or lesser degree. In 1984, the Ash Meadows population was listed as endangered under the federal Endangered Species Act of 1973. The Center for Biological Diversity (2020) has petitioned to have all Speckled Dace in the Death Valley and Owens Valley region be included under that listing, albeit with a separate listing for Long Valley Speckled Dace. The main populations of Speckled Dace in the Owens Valley are in highly modified habitats in North Fork Bishop Creek and Lower Horton, Pine, and Rock creeks, and associated ditches near Bishop (N. Buckmaster, unpublished observations). Their need for special protection is illustrated by recent disappearance of the isolated Benton Valley population of Owens Speckled Dace (N. Buckmaster unpublished observation, 2022). While captive populations are maintained by the California Department of Fish and Wildlife (CDFW), it is a tenuous situation for the long-term survival of these dace. Long Valley, Amargosa River, and Owens Valley Speckled Dace are all listed as Species of Special Concern by CDFW (Moyle et al. 2015). The causes of endangerment of dace are documented in Moyle et al. (2015) and in Center for Biological Diversity (CBD 2020)., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 521-523, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Gilbert, C. H. (1893) Report on the fishes of the Death Valley expedition collected in southern California and Nevada in 1891 with descriptions of new species. U. S. Department of Agriculture, North American Fauna, 7, 229 - 234. https: // doi. org / 10.3996 / nafa. 7.0003","Scoppettone, G. G., Hereford, M. E., Rissler, P. H., Johnson, D. M. & Salgado, J. A. (2011) Relative Abundance and Distribution of Fishes within an Established Area of Critical Environmental Concern, Inyo and San Bernardino Counties, California. Open- File Report 2011 - 1161. U. S. Department of the Interior, U. S. Geological Survey, Reston, Virginia. [unknown pagination] https: // doi. org / 10.3133 / ofr 20111161","Sada, D. W., Britten, H. B. & Brussard, P. F. (1995) Desert aquatic ecosystems and the genetic and morphological diversity of Death Valley system Speckled Dace. In: Nielsen, J. L. (Ed.), Evolution and the Aquatic System. American Fisheries Society Symposium, 17, pp. 350 - 359.","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Snyder, J. O. (1917) An account of some fishes from Owens River, California. Proceedings, United States National Museum, 54, 201 - 205. https: // doi. org / 10.5479 / si. 00963801.2233.201","Sada, D. W., Britten, H. B. & Brussard, P. F. (1993) Morphometric and genetic differentiation among Death Valley System Rhinichthys osculus. CDFG Inland Fisheries Repor t, Contract, FG 0524, 1 - 52.","Oakey, D. D., Douglas, M. E. & Douglas, M. R. (2004) Small fish in a large landscape: diversification of Rhinichthys osculus (Cyprinidae) in western North America, Copeia, 2004 (2), 207 - 221. https: // doi. org / 10.1643 / CG- 02 - 264 R 1","Mussmann, S. M., Douglas, M. R., Oakey, D. D. & Douglas, M. E. (2020) Defining relictual biodiversity: conservation units in Speckled Dace (Leuciscidae: Rhinichthys osculus) of the Greater Death Valley Ecosystem. Ecology and Evolution, 10, 10798 - 10817. https: // doi. org / 10.1002 / ece 3.6736","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Moyle, P. B. (1976) Inland Fishes of California. University of California Press, Berkeley, California, 405 pp.","Snyder, J. O. (1918) The fishes of the Lahontan system of Nevada and northeastern California. United States Bureau of Fisheries Bulletin, 35, 33 - 86.","Miller, R. R. (1973) Two new fishes, Gila bicolor snyderi and Catostomus fumeiventris from the Owens River Basin California. Occasional Papers of the Museum of Zoology, University of Michigan, 667, 1 - 19.","Moyle, P. B., Quinones, R. M., Katz, J. V. E, & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd Edition. Sacramento, California Department of Fish and Wildlife. Available from: https: // wildlife. ca. gov / Conservation / SSC / Fishes (accessed 8 February 2023)","Chen, Y., Parmenter, S. & May, B. (2007) Introgression between Lahontan and endangered Owens Tui Chubs, and apparent discovery of a new Tui Chub in the Owens Valley, California. Conservation Genetics, 8 (1), 221 - 238. https: // doi. org / 10.1007 / s 10592 - 006 - 9164 - 3","Sada, D. W. (1989) Status, distribution, and morphological variation of Speckled Dace in the Owens River system. Final Report. California Department of Fish and Game, Sacramento, California, 37 pp.","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Center for Biological Diversity (2020) Petition to List Three Populations of Speckled Dace (Rhinichthys osculus) in the Death Valley Region under the Endangered Species Act: Amargosa Canyon Speckled Dace, Long Valley Speckled Dace, Owens Speckled Dace. Submitted to US Fish and Wildlife Service, 8 June 2020, 1 - 48."]}

Rhinichthys nevadensis Gilbert 1893, new common name, Desert Speckled Dace Rhinichthys (Apocope) nevadensis, Gilbert 1893:230 Agosia nevadensis Jordan and Evermann 1896:310 Agosia nubila carringtoni, Jordan and Evermann 1896:311 Agosia robusta Rutter 1903:148; Snyder 1917:2002 Agosia carringtoni: Snyder 1913:70 Apocope robusta Snyder 1918:33; Evermann and Clark 1931:55 Rhinichthys osculus robustus: Shapovalov and Dill 1950:387; Kimsey and Fisk 1960:46; Hubbs et al. 1979:12; Moyle 2002:161. Rhinichthys osculus nevadensis Shapovalov and Dill 1950: 387; Shapovalov et al. 1981: 25; Hubbs et al 1979:12; Smith et al, 2002: 33; Moyle 2002:161; Mussmann et al. 2020:10805 Holotype: Rhinichthys (Apocope) nevadensis, USNM 46111, 65 mm SL, Ash Meadows, Nye County, Nevada, Amargosa River watershed, March 4,189 1, T. S. Palmer. Paratypes: USNM 350556, 3 (42–51 mm SL), March 4,189 1, T. S. Palmer; and USNM 46112, 5, March 3,189 1, A.K. Fisher, both from same locality as holotype. Diagnosis. Desert Speckled Dace are best recognized as small (adult lengths of 5–9 cm) cyprinoid fish that have the typical Speckled Dace phenotypic characters, inhabit Lahontan, Owens Valley and Death Valley drainages and are a distinct genetic lineage that encompasses three sublineages. The mouth is slightly subterminal, often with a frenum. Meristics and morphometrics are in the ranges shown in Table 3 but are not distinctive. Color is variable but there are usually speckles and blotches on the body which join to form a black lateral band on each side. The band is most prominent underneath each eye, sometimes extending to the snout. We recognize three subspecies, based on genomics and distribution: R. n. nevadensis, R. n. caldera, and R. n. robustus. Each has a separate account. Description. The complete original description of R. nevadensis is as follows, from Gilbert (1893: 230) although numerical fractions in the original description have been converted to decimals. All morphometric measurements are standardized by dividing into standard length. “Differing from other known species in the large head, short deep body, very small eye, and the reduction of the outer ventral ray to a mere rudiment. Head, 3.66 [times into] length (varying from 3.5 to 4.0); depth 3.66 (varying from 3.5 to 4.0); D. [dorsal rays],8; A. [anal rays],7; Lat. l. [Lateral line scales] 65. Ventrals apparently with seven rays, the outer one rudimentary, and often detected with difficulty.” “Body robust, with broad heavy head, the least depth of caudal peduncle less than half the greatest height of body. Greatest depth of head at occiput 5 in length of body. Eye very small, half the interorbital width, which equals distance from tip of snout to middle of eye, and is contained 2.66 times in head.” “Mouth terminal, very oblique, the lower jaw included, the premaxillaries not at all overlapped by the snout. The maxillary reaches the vertical from front of eye and is one-third length of head. Maxillary barbles [sic] well developed.” “Scales very irregularly placed and difficult to enumerate. The lateral line is incomplete in adults and usually does not reach to opposite dorsal fin. In the young it is variously developed, often extending though with many interruptions, to end of dorsal or base of caudal. Pores in lateral line (when complete) 58, about 66.” “Fins small, the pectorals not reaching ventrals, the latter not to vent. Front of dorsal midway between base of caudal and middle of occiput.” “In spirits, the upper half of sides is speckled and marbled with brown and the belly and lower half of sides immaculate or sparsely spotted. A broad dark lateral stripe usually present, becoming more conspicuous posteriorly, and ending in an obscure black spot on base of tail. A dark stripe sometimes present along middle of lower half of sides. ” This description is for dace now treated as a population of Amargosa Speckled Dace (R. nevadensis nevadensis) from Ash Meadows in Death Valley. It also fits the other two subspecies as well, although R. n. robustus is typically more darkly pigmented and R. n. caldera is typically pale without much dark pigmentation. Sada et al. (1993) compared meristics (Table 3) and morphometrics of dace of all three subspecies and found only small, overlapping differences, which could not be used to distinguish the three groups for identification purposes Distribution. Desert Speckled Dace are widespread in the southern Lahontan Basin and also inhabit small desert springs and streams in the Owens Valley and Death Valley regions. The common genetic heritage (ancestry) of the three subspecies is the result of late Pleistocene aquatic connections, which allowed the ancestral population(s) to spread throughout the Great Basin and into Death and Owens valleys. During the periods of high rainfall, large inter-connected lakes were formed throughout the Great Basin (Grayson 1993). One of the largest lakes was Lake Lahontan. Its remnant lakes and tributaries support Lahontan Speckled Dace to this day. Lake Russell (whose remnant is fishless Mono Lake), drained into Lake Lahontan’s southern end. About 1.3 million years ago, this connection to Lake Lahontan was blocked by vulcanism and Lake Russell subsequently spilled multiple times directly into the Owens River basin (Reheis et al. 2002). The subsequent eruption of the Long Valley caldera (0.76 mya) formed Long Valley Lake in the upper Owens Basin. Long Valley Lake, and adjacent Adobe Lake, drained into the Owens River which flowed into Owens Lake. Owens Lake spilled into Searles Lake, which drained into the Panamint Valley and Lake Manley, the site of present-day Death Valley (Grayson 1993). Lake Manly was the ultimate end for the water. Speckled Dace managed to make it through this chain, leaving populations behind in spring systems when dry desert conditions returned. The reverse movement of fish from Lake Manly to the Owens Basin was apparently geologically improbable, given the large waterfalls in the Owens River gorge and elsewhere (Hildreth & Fierstein 2016). The ancestral dace also colonized the Amargosa River which flowed into Lake Manly from the east by moving upstream from the lake. The Eastern California Lake Chain, described above, was possible because of the elevational gradient starting at about 2200 m in elevation in the Lake Russell and dropping to sea level in Lake Manly (Orme 2008). The flow, however, was not continuous because connections between lakes were variable through time. Thus, based on estimates of the timing of lake spills, Lahontan fishes were isolated in the Lake Russell Basin about 1.3 million years ago. This basin spilled numerous times. The last spill was as recently as 100,000 years ago (Reheis 2002), when the Owens River carried Lahontan fishes to Owens Lake and allowed Speckled Dace and other fishes to colonize streams flowing into the caldera. The last spill event from Owens Lake seems to have been 10–12,000 years ago (Orme 2008), although the dace had presumably already colonized Lake Manly and the Amargosa River by that time. In the Owens Valley, Long Valley Speckled Dace and the Owens Valley populations of Amargosa Speckled Dace now occupy a few spring systems or artificial refuges. They have been extirpated from most of their historic ranges. Amargosa Desert Speckled Dace apparently were once found throughout the lower elevation reaches of the Owens River and tributaries (Snyder 1917). Genetics/Genomics. The genomics-based analysis of Su et al. (2022) supports treating the Desert Speckled Dace as a widely distributed lineage, with three sub-lineages. The major lineage fits our definition of a species. The three sub-lineages can be regarded as subspecies: R. n. nevadensis, R. n. caldera, and R. n. robustus (described below). The analysis also shows that Speckled Dace from the Death Valley and Owens Valley regions share a recent ancestry (via the Amargosa River) but also show some divergence. They are best treated as Distinct Population Segments of R. nevadensis nevadensis. Notes. This analysis supports the finding that the Desert Speckled Dace is a species with three geographically and genomically defined subspecies. However, most of its distribution is encompassed by R. n. robustus in the Lahontan Basin. See subspecies accounts for more details. Etymology. Desert Speckled Dace is the new common name we propose for the Speckled Dace species which occurs in the Lahontan Basin and in the Owens and Death Valley systems. These regions are all very arid, and include Death Valley, one of the hottest and driest places on Earth. The first description of this species was based on specimens from Ash Meadows, Nevada, hence nevadensis. Genomics allows us to recognize three subspecies of Desert Speckled Dace that have been isolated from one another, Lahontan Speckled Dace, Amargosa Speckled Dace, and Long Valley Speckled Dace. Both the Lahontan Speckled Dace and the Amargosa Speckled Dace were originally described as species, R. robustus and R. nevadensis, respectively. However, they both also have a long history of being treated as subspecies of R. osculus. R. nevadensis is used as the species name for Desert Speckled Dace because it was described in 1893, nine years before R. robustus was described. The rules of zoological nomenclature declare that the oldest name used in a species description has precedence as the name for the species (International Code of Zoological Nomenclature, 1999, 4 th Edition, Chapter 11, Article 50.4. 1999), Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 517-519, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Gilbert, C. H. (1893) Report on the fishes of the Death Valley expedition collected in southern California and Nevada in 1891 with descriptions of new species. U. S. Department of Agriculture, North American Fauna, 7, 229 - 234. https: // doi. org / 10.3996 / nafa. 7.0003","Jordan, D. S. & Evermann, B. W. (1896) The Fishes of North and Middle America. Bulletin, US National Museum, 47, Part 1, 1 - 1240.","Rutter, C. (1903) Notes on fishes from streams and lakes of northeastern California not tributary to the Sacramento basin. United States Fish Commission Bulletin, 22, 145 - 148. [1902]","Snyder, J. O. (1917) An account of some fishes from Owens River, California. Proceedings, United States National Museum, 54, 201 - 205. https: // doi. org / 10.5479 / si. 00963801.2233.201","Snyder, J. O. (1913) The fishes of the streams tributary to San Francisco Bay. Bulletin of the United States Bureau of Fisheries, 32, 49 - 72.","Snyder, J. O. (1918) The fishes of the Lahontan system of Nevada and northeastern California. United States Bureau of Fisheries Bulletin, 35, 33 - 86.","Evermann, B. W. & Clark, H. W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1 - 67.","Shapovalov, L. & Dill, W. A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382 - 391","Kimsey, J. B. & Fisk, L. O. (1960) Keys to the freshwater and anadromous fishes of California. California Fish and Game, 46, 453 - 479.","Hubbs, C. L., Follett, W. I. & Dempster, L. J. (1979) List of the fishes of California. Occasional Papers of the California Academy of Sciences, 133, 1 - 54.","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Shapovalov, L., Cordone, A. J. & Dill, W. A. (1981) A list of the freshwater and anadromous fishes of California. California Fish and Game, 67, 4 - 38.","Smith, G. R., Dowling, T. E., Gobalet, K. T., Lugaski, T., Shiozawa, D. & Evans, P. (2002) Biogeography and timing rates of evolutionary events among Great Basin fishes. In: Hershler, R., Madsen, D. B. & Curry, D. R. (Eds.), Great Basin Aquatic Systems History, Smithsonian Contributions to the Earth Sciences, 33, pp. 175 - 233.","Mussmann, S. M., Douglas, M. R., Oakey, D. D. & Douglas, M. E. (2020) Defining relictual biodiversity: conservation units in Speckled Dace (Leuciscidae: Rhinichthys osculus) of the Greater Death Valley Ecosystem. Ecology and Evolution, 10, 10798 - 10817. https: // doi. org / 10.1002 / ece 3.6736","Sada, D. W., Britten, H. B. & Brussard, P. F. (1993) Morphometric and genetic differentiation among Death Valley System Rhinichthys osculus. CDFG Inland Fisheries Repor t, Contract, FG 0524, 1 - 52.","Grayson, D. K. (1993) The Desert's Past: A Natural Prehistory of the Great Basin. Smithsonian Institution Press, Washington, xx + 356 pp.","Reheis, M. C., Stine, S. & Sarna-Wojcicki, A. M. (2002) Drainage reversals in Mono Basin during the late Pliocene and Pleistocene. Geological Society of America Bulletin, 114 (8), 991 - 1006. https: // doi. org / 10.1130 / 0016 - 7606 (2002) 114 % 3 C 0991: DRIMBD % 3 E 2.0. CO; 2","Hildreth, W. & Fierstein, J. (2016) Long Valley Caldera Lake and Reincision of Owens River Gorge. U. S. Geological Survey Scientific Investigations Report, 5120, 1 - 63. https: // doi. org / 10.3133 / sir 20165120","Orme, A. R. (2008) Pleistocene pluvial lakes of the American West: a short history of research. Geological Society, London, Special Publications, 301 (1), 51 - 78. https: // doi. org / 10.1144 / SP 301.4","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388"]}

Rhinichthys klamathensis (Evermann and Meek 1898), Western Speckled Dace Fig. 6. Agosia klamathensis Evermann and Meek 1898:74; Snyder 1908:98 Apocope klamathensis Evermann and Clark 1931:55 Rhinichthys osculus klamathensis Shapovalov and Dill 1950:386; Hubbs et al. 1979:12; Shapovalov et al. 1981:25; Moyle 2002:161; Markle 2016:50 Holotype: USNM 48225 (64 mm SL) as Agosia klamathensis Evermann and Meek 1898. November 3, 1896, mouth of small creek flowing into Pelican Bay, Upper Klamath Lake, Oregon, USA, Seth Meek and A. R. Alexander. Paratypes. USNM 48231 (39 specimens) and 125012 (38 specimens). Same date and place as holotype. Diagnosis. A cryptic species readily recognized as Speckled Dace by small adult size (6–11 cm SL) and distinctive body shape. Dark blotches on sides merge into a line of dark pigmentation below eye, to snout. Meristics are in Table 3. The best way to characterize this abundant fish is as a distinct Speckled Dace lineage found throughout the Klamath, Sacramento, and Warner watersheds with three sub-lineages that cannot be readily distinguished from one another. Description. According to Evermann and Meek (1898:74-75, Fig. 6) “ the fish from [Upper Klamath Lake]… possess certain characters which serve to distinguish it from A. nubila. While the differences are slight, they are plainly evident and must be recognized, and we therefore describe it as a new species (p. 74). Head 4 [times into standard length]; depth 4.5; eye 4.5 in head; D[orsal rays]i.8; A [anal rays] i,7; [lateral line] scales about 14–78–10 (average, 73 in 49 specimens).” Body robust, subcylindrical, back somewhat elevated; snout rather long, mouth inferior, little oblique, the lower jaw included; maxillary not reaching front of orbit; upper lip without frenum; barbels present but small. Lateral line incomplete, interrupted in many places, about 30 pores developed. Origin of dorsal fin midway between front of pupil and base of caudal fin; pectorals rather short, reaching about three-fourth the distance to ventrals; ventrals reaching vent; anal[fin] large, its longest ray 1.5 in head. Color in alcohol, olivaceous, mottled and blotched with darker on back and sides; under parts pale; an obscure pale streak from eye to base of caudal fin, below which is a broad dark band; dorsals, pectorals, caudal dusky; other fins plain; a black blotch at base of caudal. This form is distinguished from Agosia nubila chiefly by the smaller size of the scales. A. nubila has 49–66 lateral line scales (mean 57, n=73), while A. klamathensis has 68–78 (mean 73, n =49). Overall, there are only minor (statistical) differences among the three subspecies from this basic description. However, Markle (2016) used lateral line scale number as the principal character to separate Western Speckled Dace from other dace taxa in Oregon. In contrast, Smith et al. (2017) could not readily distinguish lineages despite multivariate analyses of combined meristic, morphological, mtDNA, and fossil data from the entire region in which Speckled Dace sensu lato occur. Distribution. The Western Speckled Dace is found throughout the Klamath and Sacramento watersheds, including streams as far south as the Santa Maria River in southern California. Its distribution in the Warner basin is mostly in springs and small cool-water streams. Only the headwaters of Twelvemile Creek are in California (Modoc County). Before the introduction of non-native game fishes, it probably had a wider distribution within the basin. See the subspecies accounts for more detailed distributional information. Note: Geology. The Klamath River has not been connected to the ancestral Sacramento River system since the end of the Pliocene (ca. three mya); however, extensive deformation (e.g. down-faulting of the Klamath Graben) and vulcanism (both from the Cascade-arc and Medicine Lake volcanic fields) occurred almost continuously in the northern Sacramento and southern Klamath Basins (Colman et al. 2004). It is probable that this activity led to repeated drainage captures of between the two basins and allowed for intermittent gene flow among dace populations. Inter-basin connectivity presumably was less frequent between Klamath/Sacramento streams and those in the Warner Basin. The development of the Great Basin faulting and extension belt of down-faulting resulted in creation of the Warner Range (ca. 3 mya) which resulted in a permanent topographic separation of the basins sometime in the late-Pleistocene (1.0–0.1 mya) (Egger et al. 2011). As the result of these active geologic processes, each of the three major subspecies of R. klamathensis, as discussed here, is endemic to a single large watershed, plus some coastal watersheds in the case of Sacramento Speckled Dace. They share these waters with other endemic fishes. Dace ultimately colonized these three watersheds from the ancient Columbia River Basin during a series of volcanic and other geologic events that caused the predecessor of the upper Snake River to flow into the Pacific Ocean via the Klamath-Sacramento region, apparently during the Pliocene period. Fish fossils from ancient deposits in the Idaho region, that predate the rise of the Sierra Nevada and coastal mountain ranges, indicate a source of the predecessors of the highly distinctive endemic fish fauna of the lower Sacramento-San Joaquin Watershed (Minckley et al. 1986). It is likely that the Speckled Dace was not part of those early colonization events but arrived later, during a period when volcanic activity caused the upper Pit River to switch from flowing into the Sacramento River to flowing into the Klamath instead, which had its own endemic fish fauna. Some Klamath fishes (e.g. Cottus spp., Siphatales bicolor) were able to colonize the upper Pit watershed as a result. The ancestral Pit River then broke through the volcanic dike barrier and rejoined the Sacramento River system, bringing in its water members of the Klamath fish fauna. Alternately, Smith et al. (2017) suggest colonization of the Sacramento system could have come from a pass through the southern Sierra Nevada that apparently was open 3.2 million years ago or earlier. However, the genomic similarities between Sacramento and Klamath daces and between them and Warner Speckled Dace are presumably the result of their ancestors colonizing the region during a long geologically active period in the Klamath-Pit region. The genomic study of Su et al. (2022) supports this view (Figure 2) and that the Western Speckled Dace merits full species status. Note: History. The three lineages that make up this species have different taxonomic histories (Evermann and Clark 1931, Moyle 2002). The Klamath Speckled Dace was recognized in 1898 as a species, and it retained its identity even when subsumed as a subspecies under R. osculus. The Sacramento Speckled Dace was largely ignored as a taxon despite its abundance and wide distribution. Presumably early workers such as Cloudsley Rutter, Charles Gilbert, and John Otterbein Snyder saw that it could not be readily distinguished from other Speckled Dace and accepted by default the use of an improper name (some variant of Agosia carringtoni) or else treated it as Lahontan Speckled Dace. The Warner Speckled Dace came to our attention because of controversy over the status of the population in remote Foskett Spring, which modern genetic analysis showed to be part of the Warner Speckled Dace lineage described in this paper. Su et al. (2022) show that, based on genomics, the three lineages are similar enough so that together they can be treated as a single species, Rhinichthys klamathensis. Nevertheless, all three populations are isolated from one another, with long independent histories. The early history of each basin includes one or more periods of ancient interconnectedness which allowed Speckled Dace to colonize all three basins. The three lineages co-occur with other endemic fishes in their respective basins. The endemic fishes in each basin have close relatives in the other basins (e.g. Siphatales bicolor subspp.), as well as species unique to each basin. Etymology. The name ‘Western Speckled Dace’ (Markle 2016, Smith et al. 2017) covers three lineages: Klamath Speckled Dace (R. k. klamathensis), Sacramento Speckled Dace (R. k. achomawi) and Warner Speckled Dace (R. k. goyatoka). Only the Klamath Speckled Dace has been previously named, so R. klamathensis becomes the species epithet for three subspecies. Klamath is an anglicized version of a name for the native peoples inhabiting the upper Klamath Lake region in Oregon (Gudde and Bright 1998). See subspecies accounts for further explanations. Conservation Status. Sacramento and Klamath Speckled Dace are wide-spread and abundant in the large basins they inhabit. Warner Speckled Dace, including the population in Foskett Spring, are limited by the low availability of water in the desert Warner Basin. Their populations for now seem reasonably secure, although the basin has been invaded by non-native species. The Foskett Spring population was listed in 1985 as a Threatened species under the federal Endangered Species Act but it was delisted in 2019, following implementation of the recovery plan., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 525-526, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Evermann, B. W. & Meek, S. B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U. S. Fish Commission, 17, 15 - 84. https: // doi. org / 10.5962 / bhl. title. 35601","Snyder, J. O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U. S. Bureau of Fisheries Bulletin, 27, 69 - 102.","Evermann, B. W. & Clark, H. W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1 - 67.","Shapovalov, L. & Dill, W. A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382 - 391","Hubbs, C. L., Follett, W. I. & Dempster, L. J. (1979) List of the fishes of California. Occasional Papers of the California Academy of Sciences, 133, 1 - 54.","Shapovalov, L., Cordone, A. J. & Dill, W. A. (1981) A list of the freshwater and anadromous fishes of California. California Fish and Game, 67, 4 - 38.","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Markle, D. F. (2016) A Guide to the Freshwater Fishes of Oregon. Oregon State University Press, Corvallis, Oregon, 140 pp.","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Colman, S. M., Platt Bradbury, J. & Rosenbaum, J. G. (2004) Paleolimnology and paleoclimate studies in Upper Klamath Lake, Oregon. Journal of Paleolimnology, 31 (2), 129 - 138. https: // doi. org / 10.1023 / B: JOPL. 0000019235.72107.92","Egger, A. E. & Miller, E. L. (2011) Evolution of the northwestern margin of the Basin and Range: The geology and extensional history of the Warner Range and environs, northeastern California. Geosphere, 7 (3), 756 - 773. https: // doi. org / 10.1130 / GES 00620.1","Minckley, W. L., Hendrickson, D. A. & Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. & Wiley, E. O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614.","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Gudde, E. G. & Bright, W. (1998) California Place Names. University of California Press, Berkeley, California, xxviii + 467 pp. https: // doi. org / 10.1525 / 9780520920545"]}

Rhinichthys nevadensis caldera, new subspecies, Long Valley Speckled Dace Table 3, Fig. 4. Synonymy: Rhinichthys osculus subsp. Sada et al. 1995:356; Moyle 2002:161. Otherwise see synonyms for Desert Speckled Dace, Rhinichthys nevadensis. Holotype: WFB 5000 (Fig. 4). Standard length 60 mm, fork length 70 mm; White Mountain Research Center, Northeast Pond 37.360618°N 118.329370°W. Inyo County, California, July 29, 2021. Nicolas Buckmaster, Rosa Cox, California Department of Fish and Wildlife Paratypes: WFB 5001-5009 (n=9). Same as holotype. Meristic s: holotype (paratypes) Lateral line scales: 64 (59–80). Lateral line incomplete. Scales above lateral line: 10 (9–12), scales difficult to discern. Scales below lateral line: 9 (6–10), scales hard to discern Dorsal-fin rays: 8(7–8), counts include single unbranched ray. Anal-fin rays: 7 (6–7), counts include single unbranched ray. Pectoral-fin rays 13 (11–12), counts include anterior and posterior single unbranched rays. Pelvic 7 (6–8), counts include single unbranched ray. Caudal-fin rays 19 (19). Diagnosis. The Long Valley Speckled Dace is a cryptic taxon, very similar in its meristics to the Amargosa Speckled Dace (Table 3). They are distinguished from other Speckled Dace by their being endemic to the Long Valley region and having a distinct evolutionary lineage, as revealed by genomics (Su et al. 2022). Adults are small (rarely more than 75 mm SL) and recognizable as Speckled Dace by their thick caudal peduncle, sub-cylindrical body, small fins, small eyes (relative to head), and blunt, pointed snout. In life, they are a bronzy color with light speckling (Fig. 6). Striping is largely absent. Maxillary barbels and a frenum are usually absent. They can be distinguished statistically from similar Amargosa Speckled Dace by slightly higher average numbers of pectoral and pelvic, higher lateral line scale count, lower lateral line pore scale count, and absence of maxillary barbels (Sada et al. 1989, 1995). The following mean counts (standard error) are from Long Valley Speckled Dace collected in Whitmore Hot Springs and at an unnamed spring at Little Alkali Lake (Sada 1989, Table 3): lateral line scales 61.7 (1.4); lateral line scales with pores 19.0 (5.0); dorsal-fin rays 8.0 (0.0); anal-fin rays 7.0 (0.0); pectoral-fin rays 13.0 (0.4); pelvic 7.4 (0.2). Description. Long Valley Speckled Dace are small, active fish with the typical Speckled Dace morphology, as described for the species. Genetics/Genomics. Sada et al. (1989, 1995), using isozymes and morphometrics, were the first to recognize Long Valley Speckled Dace as different from other dace populations in the region. Oakey et al. (2004), Mussmann et al (2020) and Su et al (2022) all found genetic differences that separated Long Valley Speckled Dace from Amargosa and Lahontan Speckled Dace. Distribution. The historic range of this dace was in the outlets of hot springs and associated marshes in the remains of the Long Valley volcanic caldera, just east of Mammoth Lakes, Mono County, as well as in Hot Creek. It quite likely had its origins when Speckled Dace colonized the upper Owens Valley region from the Mono Lake Basin via Adobe Valley, during a late Pleistocene pluvial period. During this time, Mono Lake levels were high enough so that it periodically spilled into Adobe Valley, which drained into the Owens River, from which fish presumably moved upstream into Long Valley. Subsequent down-faulting of the Owens River and formation of steep waterfalls in the Owens River gorge likely isolated Long Valley from the Owens Valley around 100,000 years ago (Hildreth and Fierstein 2016). Note. Long Valley Speckled Dace were considered to be one of the scattered populations of Owens Valley Speckled Dace until Sada et al. (1993, 1995) showed it was genetically and morphometrically distinct and that it was closely related to dace populations in Ash Meadows, Amargosa River, and elsewhere in Owens Valley and Death Valley. Moyle (2002) and Moyle et al. (2015) recognized it as an undescribed subspecies. Our analysis shows that the Long Valley Speckled Dace is a distinct lineage confined to a small part of the Owens Valley region. It merits subspecies designation based on the following lines of evidence. Taxonomy. Multiple analyses indicate that Long Valley dace is a distinct lineage (subspecies) within Desert Speckled Dace, R. nevadensis (Oakey et al 2004, Mussmann et al. 2020, Sada et al. 1995, Moyle 2002, Moyle et al. 2015 and Su et al., 2022). Geography/geology. During pluvial periods of the Pleistocene, the Death Valley region was a series of interconnected large lakes, with abundant fishes (Hubbs and Miller 1948). When the climate changed and heavy precipitation stopped falling, the lakes and rivers dried up or became small remnants of what they once were. This process resulted in numerous isolated Speckled Dace populations with little, or at least very infrequent, opportunity for genetic exchange. See Desert Speckled Dace account for more details. Long Valley is part of the Death Valley regional endemism hotspot with numerous endemic plants and animals, including fishes (Sada et al. 1995). Long Valley Speckled Dace can be regarded as another organism endemic to the isolated Owens Valley region. Genetics/genomics. Sada et al. (1993, 1995) were first to recognize that Long Valley Speckled Dace were genetically distinct from other dace, a finding confirmed by other studies (Mussmann et al. 2020, Su et al. 2022). See Amargosa Speckled Dace (R. nevadensis nevadensis) description for more details. Etymology. The Long Valley Speckled Dace was historically known only from small streams flowing into the Long Valley Caldera, the remnants of a gigantic volcano that last erupted 0.7 million years ago (Hildreth and Fierstein 2016). They colonized the remnants of the caldera during the late Pleistocene, hence caldera. Conservation Status. Long Valley Speckled Dace have been extirpated from all but one of their historic collection sites, including Hot Creek. The only population left in its native range (as of 2021) is in Whitmore Marsh and its inlet stream in Long Valley (Moyle et al. 2015). Unfortunately, this marsh is now maintained by the outflows of a hot spring system that has been developed as a public swimming pool by the Town of Mammoth Lakes. Discharge is approximately 2 cfs and is lightly chlorinated. The outlet stream feeds an alkaline marsh of roughly 1 acre. In 1989, dace occupied 250 meters of stream and two large shallow ponds less than a half meter deep (Moyle et al 2015). Surveys in 2002 and 2009 by CDFW found this population to be relatively stable (S. Parmenter, CDFW, pers. comm. 2009). In 2018, 2019, and 2020, they were not found in these habitats (S. Parmenter, CDFW, in Center for Biological Diversity, 2020). However, in 2021 and 2022, trapping produced a few fish (R. Black, CDFW, pers. comm, 2021). The only other population is in a single artificial pond into which they were introduced at the White Mountain Research Center, outside their native range; the status of this dace population is checked by CDFW on a routine basis (R. Black, CDFW, pers. comm.). The multiple causes of decline are discussed in Moyle et al. (2015) and Center for Biological Diversity (2020). The CBD (2020) has petitioned to have this dace listed as endangered under the federal ESA or else have all Speckled Dace in the Death Valley region included under the 1984 listing of Ash Meadows Speckled Dace as endangered. It is listed as a Species of Special Concern (critical concern) by CDFW., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 523-524, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Sada, D. W., Britten, H. B. & Brussard, P. F. (1995) Desert aquatic ecosystems and the genetic and morphological diversity of Death Valley system Speckled Dace. In: Nielsen, J. L. (Ed.), Evolution and the Aquatic System. American Fisheries Society Symposium, 17, pp. 350 - 359.","Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Sada, D. W. (1989) Status, distribution, and morphological variation of Speckled Dace in the Owens River system. Final Report. California Department of Fish and Game, Sacramento, California, 37 pp.","Oakey, D. D., Douglas, M. E. & Douglas, M. R. (2004) Small fish in a large landscape: diversification of Rhinichthys osculus (Cyprinidae) in western North America, Copeia, 2004 (2), 207 - 221. https: // doi. org / 10.1643 / CG- 02 - 264 R 1","Mussmann, S. M., Douglas, M. R., Oakey, D. D. & Douglas, M. E. (2020) Defining relictual biodiversity: conservation units in Speckled Dace (Leuciscidae: Rhinichthys osculus) of the Greater Death Valley Ecosystem. Ecology and Evolution, 10, 10798 - 10817. https: // doi. org / 10.1002 / ece 3.6736","Hildreth, W. & Fierstein, J. (2016) Long Valley Caldera Lake and Reincision of Owens River Gorge. U. S. Geological Survey Scientific Investigations Report, 5120, 1 - 63. https: // doi. org / 10.3133 / sir 20165120","Sada, D. W., Britten, H. B. & Brussard, P. F. (1993) Morphometric and genetic differentiation among Death Valley System Rhinichthys osculus. CDFG Inland Fisheries Repor t, Contract, FG 0524, 1 - 52.","Moyle, P. B., Quinones, R. M., Katz, J. V. E, & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd Edition. Sacramento, California Department of Fish and Wildlife. Available from: https: // wildlife. ca. gov / Conservation / SSC / Fishes (accessed 8 February 2023)","Center for Biological Diversity (2020) Petition to List Three Populations of Speckled Dace (Rhinichthys osculus) in the Death Valley Region under the Endangered Species Act: Amargosa Canyon Speckled Dace, Long Valley Speckled Dace, Owens Speckled Dace. Submitted to US Fish and Wildlife Service, 8 June 2020, 1 - 48."]}

Rhinichthys klamathensis achomawi, new subspecies. Sacramento Speckled Dace. Figs. 1, 4D, Synonymy. See R. klamathensis account for subspecies synonymies. Holotype: WFB-3171 (Figure 1), 70 mm SL. Bear Creek at Pondosa Bridge, 1.6 km SW of Pondosa, Siskiyou County, California. 41.1884°N - 121.700667°W. July 20, 2015. Jason Baumsteiger, Mollie Ogaz, Christopher R. Jasper, Tyler R. Goodearly, and Matthew J. Young Paratypes WFB - 3171a-3171i (n=9). data the same as holotype. Meristics: holotype (paratypes) Standard length: 70 (41–56 mm) Lateral line scales: 74 (63–73). Lateral line incomplete in most; counts include 2–3 scales beyond lateral line. Scales above lateral line: 13 (11–14) Scales below lateral line: 9 (8–10) Dorsal-fin rays: 8(8), counts include single unbranched ray. Anal-fin rays: 7 (7), counts include single unbranched ray. Pectoral-fin rays 15 (12–15), counts include unbranched rays. Pelvic 7 (8–9), counts include unbranched rays. Caudal-fin rays 19 (19). Basic morphology is described below. Barbels and frenum are usually present. Diagnosis. Same as for Western Speckled Dace, R. klamathensis. Distinguished by distribution and genomics as the Speckled Dace endemic to aquatic habitats in Sacramento River drainage basin and associated coastal drainages. Description. Sacramento Speckled dace are identical to Klamath, Warner, and Lahontan Speckled Dace based on meristics and morphometrics. This is presumably the reason they have been undescribed for so long. The general description of Speckled Dace in Moyle (2002:161) was based on Sacramento dace, so fits the subspecies well: The Speckled Dace is a small (usually Distribution. The Sacramento Speckled Dace is found in streams and lakes in the Goose Lake (Oregon and California) and Pit River watersheds, the Sacramento River and its tributaries, the Salinas basin, and, in the far south, San Luis Obispo Creek, Arroyo Grande, and the Santa Maria River (Figure 3). It is conspicuous by its absence from the San Joaquin River watershed, although Rutter (1908) collected two small dace from the Kings River at Centerville. Collections made on Los Gatos Creek, an east-side tributary to the San Joaquin River, in 1941 contained Speckled Dace but there are no records of them since then (R. Leidy, pers. comm., 2021). Sacramento Speckled Dace are absent naturally from the Russian River watershed and from all other coastal watersheds, except the San Lorenzo River, Salinas River, San Luis Obispo Creek and the Santa Maria River in southern California. They are also absent from Clear Lake (Lake County) and its tributaries. However, they are present in Cache Creek which flows out of Clear Lake (Moyle 2002, unpublished data). Zoogeography. The Sacramento basin is a large, well-defined region, isolated by the Sierra Nevada Range on the east, the Coast Range on the west, and the Cascade Range in the north. The ancestors of this highly endemic freshwater dispersant fish fauna have ancient roots in ancient lakes and rivers of the region to the east, now drained mostly by tributaries to the Snake River (Minckley et al 1986; Moyle 2002). Speckled Dace presumably entered the region by first colonizing the Klamath Basin with other Great Basin fishes and then by expanding their population to the Sacramento-San Joaquin basin when Klamath River headwaters connected to the region via the Pit River. Dace then found it possible to colonize most of the entire basin (except where noted), reaching as far south as the Santa Maria River (via the Pajaro-Salinas River system). The Sacramento Speckled Dace thus became a member of one of the most highly endemic assemblages of freshwater dispersing fishes in the western USA. The Sacramento Speckled Dace, however, may have been a rather late addition to this fauna, which would explain its close genetic relationship with Klamath Speckled Dace. Genetics/genomics. The Sacramento Speckled Dace is one of three lineages, defined by genomics, that make up the Western Speckled Dace, Rhinichthys klamathensis (Smith et al. 2017; Su et al. 2022). The three lineages are also supported by Wiesenfeld et al. (2017) and Smith et al. (2017) using mtDNA and microsatellites. Etymology. The species name honors the Achomawi people (“river people”) whose historic homelands were along the Pit River and tributaries in northeastern California, from Goose Lake to Pit River Falls, and beyond (Dixon 1908). The Pit River is a branch of the Sacramento River that drains much of northeastern California. This region includes the Fall River system and Achjumawi Lava Springs State Park. The Speckled Dace is one of the most abundant native fishes in streams flowing through Achomawi lands. Alternate spellings include Achjumawi and Achumawi (Dixon 1908). The scientific names have a confusing history, starting with Jordan and Evermann (1896), who did not mention by name Speckled Dace in the Sacramento or Klamath rivers. They did include Sacramento Speckled Dace, vaguely from California, within Agosia nubila carringtoni. This taxon was first described in 1876 as Apocope carringtoni from Utah. Jordan and Evermann (1896) basically used this taxon to include miscellaneous Great Basin populations, and provisionally fish from the Lahontan Basin, as well as from various localities in southern and central California. “These California and Nevada forms may be distinct species, but if so, we are unable to define them (Jordan and Evermann 1896:312).” Snyder (1908) followed Jordan and Evermann (1896) in using A. n. carringtoni for a grab-bag of Speckled Dace from Oregon and California. By default, carringtoni became the species epithet for Sacramento Speckled Dace as Agosia carringtoni (Snyder 1913, 1917) and Rhinichthys osculus carringtoni (Shapovalov and Dill 1950, Kimsey and Fisk 1960). Moyle (1976, 2002) did not provide a name for Sacramento populations. See Table 1 for other synonyms. Conservation Status. The Sacramento Speckled Dace is widespread and abundant in the Sacramento River and tributaries, the Pit River system, Pajaro-Salinas watershed, San Luis Obispo Creek, and the Santa Maria River. It seems to have been extirpated from the San Joaquin River and tributaries although old records are scarce. The Speckled Dace is also absent from streams tributary to the San Francisco Estuary, including the Guadalupe River and tributaries, Coyote Creek, and Alameda Creek, for which there are historic records (Leidy 2007, R.L. Leidy, pers. comm. 2021). The dace was extirpated from Coyote Creek in the 1970s (Scoppettone and Smith 1976; J.J. Smith, pers. comm. 2021) and from Alameda Creek in the early 1900s (R. Leidy, pers. com. 2021). It has been extirpated from the Pajaro River, which flows into Monterey Bay, although it is still found in the upper and middle reaches of the San Benito River, a tributary to the Pajaro (J. J. Smith, pers. comm. 2021). The historic records, though few, indicate that the Speckled Dace was extirpated from these watersheds as the result of dams, habitat change, and diversions of water, coupled with severe drought., Published as part of Moyle, Peter B., Buckmaster, Nicholas & Su, Yingxin, 2023, Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA, pp. 501-539 in Zootaxa 5249 (5) on pages 529-531, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/7701357, {"references":["Moyle, P. B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.","Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin, with a study of their distribution and variation. United States Bureau of Fisheries Bulletin, 27, 105 - 152. [1907]","Cornelius, R. H. (1969) The Systematics and Zoogeography of Rhinichthys osculus (Girard) in Southern California. M. A. Thesis, California State University, Fullerton, California, 195 pp.","Sada, D. W., Britten, H. B. & Brussard, P. F. (1993) Morphometric and genetic differentiation among Death Valley System Rhinichthys osculus. CDFG Inland Fisheries Repor t, Contract, FG 0524, 1 - 52.","Minckley, W. L., Hendrickson, D. A. & Bond, C. E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C. H. & Wiley, E. O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519 - 614.","Smith, G. R., Chow, J., Unmack, P. J., Markle, D. F. & Dowling, T. E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1 - 83","Su, Y., Moyle, P. B., Campbell, M. A., Finger, A. J., O'Rourke, S., Baumsteiger, J. & Miller, M. R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695 - 710. https: // doi. org / 10.1002 / tafs. 10388","Dixon, R. B. (1908) Notes on the Achomawi and Atsugewii Indians of northern California. American Anthropologist, 10 (2), 208 - 220. https: // doi. org / 10.1525 / aa. 1908.10.2.02 a 00030","Jordan, D. S. & Evermann, B. W. (1896) The Fishes of North and Middle America. Bulletin, US National Museum, 47, Part 1, 1 - 1240.","Snyder, J. O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U. S. Bureau of Fisheries Bulletin, 27, 69 - 102.","Snyder, J. O. (1913) The fishes of the streams tributary to San Francisco Bay. Bulletin of the United States Bureau of Fisheries, 32, 49 - 72.","Snyder, J. O. (1917) An account of some fishes from Owens River, California. Proceedings, United States National Museum, 54, 201 - 205. https: // doi. org / 10.5479 / si. 00963801.2233.201","Shapovalov, L. & Dill, W. A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382 - 391","Kimsey, J. B. & Fisk, L. O. (1960) Keys to the freshwater and anadromous fishes of California. California Fish and Game, 46, 453 - 479.","Moyle, P. B. (1976) Inland Fishes of California. University of California Press, Berkeley, California, 405 pp.","Leidy, R. A. (2007) Ecology, Assemblage Structure, Distribution, and Status of Fishes in Streams Tributary to the San Francisco Estuary. San Francisco Estuary Institute Contribution, 530 pp."]}

Cottus gulosus gulosus (Girard 1854), San Joaquin Riffle Sculpin, nominate subspecies Description is the same as for the C. gulosus (Fig. 1) Holotype: USNM 291. Same as for C. gulosus, above. Paratype: None designated Diagnosis: Distinguished from other members of the C. gulosus complex as a distinct lineage as determined by genomic studies (Baumsteiger et al. 2014, this study) and by its limited geographic distribution (Fig. 3). See species description for distinguishing it from co-occurring C. asper. Distribution. These California endemic sculpins are found in cold-water riffles, in headwaters or below dams with cold water releases into rivers in the southern Sierra Nevada on the eastern side of the Central Valley, except for the American River (which flows directly into the Sacramento River), the rivers that contain now-isolated populations all flow into the southern Central Valley, especially the San Joaquin River. Rivers with populations include the Mokelumne, Stanislaus, Tuolumne, Merced, Fresno, San Joaquin, Kings, and Kern rivers. Etymology. See C. gulosus, Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on page 517, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473, {"references":["Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165.","Baumsteiger, J., Kinziger, A. P., Reid., S. B. & Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758"]}

Cottus ohlone pomo Moyle and Campbell 2022. Pomo Riffle Sculpin, new subspecies Description as for C. gulosus. Meristics of sculpins used in this study (Table 1, n = 21): anal-fin rays (12–14), dorsal spines (6–8), dorsal rays (15–19), dorsal fins joined? (variable), black spot on dorsal (yes), palatine teeth present (no), maxilla reaches eye (variable), chin pores (1–2), pelvic fins spines + rays (1+3), pelvic fins do not touch vent, pectoral -in rays (14–16), preopercular spines (2), lateral line complete, lateral line pores 27–35. Other characters are listed in Table 1. Holotype. WFB 3396 90 mm SL, 105 mm TL from Pieta Creek, upstream of mouth on Russian River, Mendocino County, California, N 38.925278 W 123.054500. Collected by J. Baumsteiger and P. Moyle, August 2, 2017. 90 mm SL, 105 mm TL. Anal-fin rays 13; dorsal-fin spines 8; dorsal-fin rays 17; dorsal fins not joined; dorsal fin spot present; palatine teeth absent; maxilla reaches middle of eye; chin pores 2; pelvic fins 1 spine, 3 rays; pelvic fins do not touch vent; pectora1-fin rays 14; preopercular spines 1; lateral line incomplete; lateral line pores 31; axillary prickles present. Paratypes. WFB 3397, 3398, 3399, 3400 from same location. 71–81 mm SL, 91–105 mm TL. Anal fin rays 13; dorsal fin spines 8; dorsal fin rays 17; dorsal fins not joined; dorsal fin spot variable in presence; palatine teeth absent; maxilla reaches middle of eye; 2 chin pores; pelvic fins 1 spine, 3 rays; pelvic fins do not touch vent; pectora1 fin rays 14–15; preopercular spines 1; lateral line completeness variable; lateral line pores, 25–30; axillary prickles present. Diagnosis. Distinguished from other members of the C. gulosus complex as a distinct lineage as determined by genomics and by its distinctive distribution (Fig. 1). Distribution. Present in the upper portions of the Russian River, above the mouth of Mark West Creek, including the East Fork, as well as in tributaries to northern San Francisco Bay, including the Napa River, Petaluma River, Sonoma Creek, and smaller tributaries (Fig. 3). These streams had connections in the past to the Russian River, via the shifting headwaters of Sonoma Creek. Etymology: Ohlone honors the Ohlone people, as discussed in the names section for C. o. ohlone (see http:// www.muwekma.org/). Pomo honors the diverse native peoples who were once the principal human inhabitants of the Russian River region, which contains streams that are important habitat for Pomo Riffle Sculpin., Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on page 519, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473

Cottus gulosus wintu Moyle and Campbell 2022. Sacramento Riffle Sculpin, new subspecies. Description as for C. gulosus (Table 1, Fig. 1). Characteristics of C. g. wintu used in this study (n=15) are as follows: anal fin rays (12–15), dorsal spines (7–10), dorsal rays (14–19), dorsal fins joined? (variable), black spot on dorsal (yes), palatine teeth present (variable), mouth vs eye (variable but maxilla mostly reaches mid-eye), chin pores (0–2), pelvic fins spines + rays (1+3), pelvics fins mostly do not touch vent, pectoral fin rays (14–16), preopercular spines (1–2), lateral line variable, lateral line pores (26–38). Other characters are listed in Table 1. Holotype. WFB 3424. 78 mm SL, 96 mm TL. Fig. 1. North Fork Feather River, Butte County, California. Coll. Jason Baumsteiger, Oct. 5, 2017. Anal-fin rays 13; dorsal-fin spines 8; dorsal-fin rays 17; dorsal fins joined; dorsal fin spot present; palatine teeth absent; maxilla reaches middle of eye; 1 chin pore; pelvic fins 1 spine, 3 rays; pelvics do not touch vent; pectora1 fin rays 14; preopercular spines 1; lateral line complete; lateral line pores, 35; axillary prickles present. Paratypes (4). WFB 3425–3428. 78–89 mm SL, 96–108 TL. Anal-fin rays 13; dorsal spines 8; dorsal-fin rays 17–18; dorsal fins joined; dorsal fin spot present on 3; palatine teeth absent; maxilla reaches middle of eye; 1–2 chin pores; pelvic fins 1 spine, 3 rays; pelvics do not touch vent; pectora1 fin rays 15; preopercular spines 1; lateral line complete; lateral line pores, 36–41; axillary prickles present. Distribution. Found in the Sacramento River watershed (northern Central Valley), from the American River (and tributaries) north to Shasta Dam. Above Shasta Reservoir, it occurs primarily in the upper Sacramento River and its tributaries (Baumsteiger et al. 2014). Other tributaries to Shasta Reservoir (e.g. McCloud River, Squaw Valley Creek) support C. pitensis, while the reservoir itself supports C. asper (Prickly Sculpin). It inhabits cool-water permanent tributaries and rivers and rivers below dams, where reservoir releases provide cool water (generally less than 24°C). Etymology. The species name honors the Wintu people who were the region’s original inhabitants, with their lands encompassing many of the streams and rivers where this subspecies lives. The Winnemem Wintu (The Middle Water People) still live in the area and are working to restore salmon runs and to protect sacred rivers and sites in the region., Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on page 517, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473, {"references":["Baumsteiger, J., Kinziger, A. P., Reid., S. B. & Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758"]}

Cottus ohlone Moyle and Campbell 2022. Coastal Riffle Sculpin, new species urn:lsid:zoobank.org:act: 8F6E51D6-5317-4F4E-9246-B2F30FBE7207 Description as for C. gulosus. Meristics of sculpins used in this study (Table 1, n = 45): anal-fin rays (12–14), dorsal spines (6–9), dorsal-fin rays (15–19), dorsal fins joined? (variable), black spot present on dorsal, palatine teeth usually present (variable), mouth vs eye (variable but maxilla mostly reaches mid-eye), chin pores (0–2), pelvic fin spines + rays (1+3), pelvics do not touch vent, pectoral-fin rays (14–16), preopercular spines (2), lateral line (complete/incomplete, variable), lateral line pores (25–35). Other characters are listed in Table 1. Holotype. WFB- 3402. 65 m SL, 79 mm TL from Guadalupe Creek, Santa Clara County, California. November 18, 1986. Collected by Jerry J. Smith (Fig. 1). 65 mm SL, 79 mm TL. Anal fin rays 13; dorsal fin spines 8; dorsal fin rays 19; dorsal fins joined; dorsal fin spot present; palatine teeth absent; maxilla reaches middle of eye; 1 chin pore; pelvic fins 1 spine, 3 rays; pelvics do not touch vent; pectora1-fin rays 15; preopercular spines 1; lateral line incomplete; lateral line pores, 27; axillary prickles present but small. Paratypes. WFB-3403, 3404, 3405, 3406 from same location. 58–60 mm SL, 74–81 TL. Anal-fin rays 13; dorsal fin spines 8; dorsal fin rays 17–19; dorsal fins variable in joining; dorsal fin spot present; palatine teeth absent; maxilla reaches middle of eye; 1–2 chin pores; pelvic fins 1 spine, 3 rays; pelvic fins do not touch vent in 3 of 4; pectora1 fin rays 13–15; preopercular spines 1; lateral line incomplete in most; lateral line pores, 26–33; axillary prickles present but small. Diagnosis. Distinguished from other members of the C. gulosus complex as a distinct lineage as determined by molecular phylogenetic and population genomic evidence (Baumsteiger et al. 2014, this study) and by its limited distribution (Fig. 2). Distribution. Restricted to drainages flowing west or southwest along the Coast Range Mountains of California, with two distinct lineages, described as subspecies here. The watersheds include (a) the Russian River and Redwood Creek (which flow directly into the Pacific Ocean, (b) the Napa River, Sonoma Creek and nearby watersheds that flow into north San Francisco Bay, (c) streams draining the Diablo and Coastal ranges through the Santa Clara Valley and nearby areas and flowing into south San Francisco Bay, and (d) tributaries to the Pajaro River, in the Salinas River watershed (Fig. 2). Populations may also exist in the headwaters of the Salinas River (Snyder 1913) although there are no recent records. Habitats are restricted to cool, clear permanent streams with rocky riffles, mostly in headwaters. Etymology. Ohlone honors the Ohlone peoples, who were the original human inhabitants of the Santa Clara Valley region and much of the southern San Francisco Bay region (see http://www.muwekma.org/). The name Ohlone refers to the more than 50 peoples that spoke similar languages and interacted with one another in the region. Their descendants are largely encompassed in the present-day Muwekma Ohlone Tribe., Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on page 518, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473, {"references":["Baumsteiger, J., Kinziger, A. P., Reid., S. B. & Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758","Snyder, J. O. (1913) The fishes of the streams tributary to San Francisco Bay. Bulletin of the United States Bureau of Fisheries, 32, 49 - 72."]}

Cottus gulosus (Girard 1854), Inland Riffle Sculpin (Fig. 1) Originally described by Charles Girard as Cottopsis gulosus from the San Joaquin River, as follows: “Largest specimens a little over three inches {76 cm} in total length; of which the head forms the third, the caudal fin excepted. Preopercle provided with two small spines, such as may be observed in several species of Cottus proper, the head otherwise is smooth; mouth proportionately large; posterior extremity of upper maxillary reaching a vertical line, which would pass behind the pupil. A space of five twentieths of an inch exists between the origin of the first dorsal and the occiput. Second dorsal connected with the first by a low membrane proceeding from the last spiny ray. Tips of posterior soft rays not quite reaching the base of caudal. Anal, about as high as second dorsal, but shorter. Caudal, well developed and posteriorly rounded. Origin of ventrals midway between the tip of lower jaw and the anus. Pectorals broad and large, its longest rays extending posteriorly as far as the fifth ray of second dorsal. D IX. 18. A 13. C 3. 1. 5. 4. 1. 2. V I. 4. P 15. Lateral line uninterrupted from thoracic region to base of caudal. Abdomen beset with minute prickles; skin elsewhere smooth. Ground color reddish brown; head and dorsal region spotted with black. Dorsal, caudal and pectorals barred with black; first dorsal provided posteriorly with an elongated black spot. Anal and ventrals unicolor. (Girard 1854; p.130). This description still fits the species complex today, although larger samples show more variation in fin ray counts and characters (Table 1). Characteristics of Riffle Sculpins used in this study (n = 23) are as follows: anal-fin rays (12–15), dorsal spines (6–10), dorsal rays (14–19), dorsal fins usually joined, black spot on dorsal (yes), palatine teeth present (variable), mouth vs eye (variable but maxilla mostly reaches mid-eye), chin pores (0–2), pelvic fins spines + rays (1+2–3), pelvics touch vent when depressed (variable but mostly not), pectoral fin rays (13–16), preopercular spines (1–2), lateral line completeness variable, and lateral line pores (26–38). Holotype. USNM 291, Cottopsis gulosus from San Joaquin River, California (two specimens), Collected by A. L. Heermann and described by Charles Girard (1854). Diagnosis. It is distinguished from other members of the C. gulosus complex as a distinct lineage determined by genomic studies and its limited geographic distribution (Figs. 1, 3). In streams, it can co-occur with C. asper from which it can be most easily separated by its short anal fin (12–15 rays vs. 16–19 for C. asper) and short pelvic fins, which do not touch the vent when depressed. Distribution. C. gulosus is the name historically assigned to all populations in the species complex as described in this paper, including C. pitensis. Our study shows that the name should be restricted to (a) populations found in streams in the western Sierra Nevada, on the east side of the southern Central Valley, from the American River watershed in the north to the Kern River watershed in the south (Fig. 2), including the San Joaquin River and its tributaries and (b) populations in the Sacramento River and streams tributary to it, except the Pit River. Baumsteiger et al. (2014) used mitochondrial and nuclear DNA to examine a large sample (n = 872) of sculpins assigned to the C. gulosus species complex. They found a distinct, genetically-based separation of C. gulosus from streams flowing into the southern Central Valley, from similar sculpins in streams flowing into the northern Central Valley (Sacramento Valley). These populations here are treated here as new subspecies. The two river systems join in the tidal Sacramento-San Joaquin Delta. A similar break was found in the distribution of the Sacramento Riffle Sculpin lineage at the mouth of the Pit River (now under Shasta Reservoir); sculpins above this break are genetically Pit Sculpins while those below the break are a subspecies that shows evidence of ancient hybridization with C. pitensis (Baumsteiger et al. 2014). Etymology. Cottus was the Roman name for European sculpin, while gulosus roughly translates as ‘big mouth’ or ‘gluttonous’ (Moyle 2002)., Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on page 516, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473, {"references":["Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165.","Baumsteiger, J., Kinziger, A. P., Reid., S. B. & Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758","Moyle, P. B. (2002) Inland Fishes of California. Revised and Expanded. University of California Press, Berkeley, 517 pp."]}

Cottus pitensis Bailey and Bond 1963. Pit Sculpin (Fig. 3). Cottus pitensis is described in detail by Bailey and Bond (1963), a description that largely overlaps that of C. gulosus. Diagnostic features are given below. Holotype. UMMZ 130558, adult male 83 mm SL, from North Fork, Pit River, 1934. Paratypes. UMMZ 130559, six specimens 55–76 mm SL. Diagnosis. “Distinguished from gulosus by consistent absence of palatine teeth, the usually better-developed lateral line, and the typical complement of two preopercular spines (Bailey and Bond 1963, p. 21). According to Bailey and Bond (1963), C. gulosus has 2–4 preopercular spines but usually 3, while C. pitensis has 1–3, but usually 2. In the original description, Girard (1854) noted that C. gulosus has two preopercular spines. All other characteristics show broad overlap (Bailey and Bond 1963). Genomic studies (Baumsteiger et al. 2014, this study) confirm its species status, as does the distinct distribution. See Table 1 for meristic counts on specimens used in this study (n=8). Distribution. Endemic to the Pit River watershed in northeastern California and southeastern Oregon. Oregon populations are found in tributaries to Goose Lake, including Drews, Cottonwood, and Thomas creeks (Lake County, Oregon). In California, the Pit Sculpin is found in tributaries to Goose Lake (Lassen and Willow Creeks) and then south in streams throughout the Pit River watershed (Modoc and Shasta counties) to Squaw Valley Creek, now a tributary to Shasta Reservoir. Kinziger et al. (2016) note that the complex geologic history of the region has resulted in some population structure in the Pit Sculpin, suggesting further investigation of its taxonomic relationships is needed. Etymology. Named after the Pit River, which encompasses its distribution. The Pit River was so named because early Euro-American visitors were impressed by the deep pits dug by the native peoples to trap wildlife (Gudde and Bright 1998)., Published as part of Moyle, Peter B. & Campbell, Matthew A., 2022, Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA, pp. 501-527 in Zootaxa 5154 (5) on pages 517-518, DOI: 10.11646/zootaxa.5154.5.1, http://zenodo.org/record/6655473, {"references":["Bailey, R. M. & Bond, C. E. (1963) Four new species of freshwater sculpins, genus Cottus, from western North America. Occasional Papers of the Museum of Zoology, University of Michigan, 634, 1 - 27.","Girard, C. (1854) Descriptions of new fishes collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U. S. A. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 129 - 165.","Baumsteiger, J., Kinziger, A. P., Reid., S. B. & Aguilar, A. (2014) Complex phylogeography and historical hybridization between sister taxa of freshwater sculpin (Cottus). Molecular Ecology, 23, 2602 - 2618. https: // doi. org / 10.1111 / mec. 12758","Kinziger, A. P., Hellmair, M., Fong, S. R., Goodman, D. H. & Kelsey, H. (2016) Evolution of rough sculpin (Cottus asperrimus): genetic divergence and late Quaternary displacement on the Hat Creek fault, California, USA. Conservation Genetics, 17 (6), 1257 - 1267. https: // doi. org / 10.1007 / s 10592 - 016 - 0859 - 9","Gudde, E. G. & Bright, W. (1998) California Place Names. University of California Press, Berkeley, 172 pp. https: // doi. org / 10.1525 / 9780520920545"]}

Moyle, Peter B., Campbell, Matthew A. (2022): Cryptic Species of Freshwater Sculpin (Cottidae: Cottus) in California, USA. Zootaxa 5154 (5): 501-527, DOI: https://doi.org/10.11646/zootaxa.5154.5.1

Hesperoleucus venustus Snyder, 1913 Coastal Roach Holotype: USNM 74476. Originally described from Coyote Creek, Santa Clara Co., CA. Paratypes: CAS-SU 22492, 24282; USNM 75335. Snyder (1913) originally split this species into three species: H. navarroensis (from the Navarro River), H. venustus (from the Russian River), and H. subditus (Monterey region). These populations were later considered to be subspecies within the single species CA Roach, as suggested by the unpublished study of Murphy (1948). Aguilar & Jones (2009) did not find strong support for these species or subspecies with their mtDNA analysis although their microsatellite analysis found some structure among regions. As mentioned above, Baumsteiger et al. (2017) found clear evidence for populations in all coastal draining watersheds belonging to a single species. Subspecies were also detected but more broadly distributed than previously thought. Watersheds extending from the Russian River north to the Eel River (excluding the Gualala River) clustered as one subspecies and watersheds south of the Russian River (Tomales Bay to the Monterey region) clustered as a second. Description. Based on previous morphological work, the Coastal Roach species has a mixture of characters (Table 1). For a cyprinid, the head is large and conical, the tail deeply forked, the eyes large, and the mouth subterminal, slanting at a downward angle. However, the body can be stout or slender, the snout short or more pointed, the caudal peduncle narrow or thick, and the fins small and rounded or long and narrow. Individuals are typically less than 120 mm total length. The dorsal fin is short (7���10 rays) and positioned behind the insertion point of the pelvic fin. The anal fin has 6���9 rays. Pharyngeal teeth (0,5���4,0) have curved tips which overhang grinding surfaces of moderate size. Gill rakers number 8���11. The scales are small, numbering 47���63 along the lateral line and 32���38 before the dorsal fin. Individuals are usually dark on the upper half of their bodies, ranging from a shadowy gray to a steel blue, while the lower half of the body is much lighter, usually a dull white/silver color. A light lateral stripe approximately two scales wide extends from upper edge of the gill opening to the base of tail and entirely above the lateral line. Below this line is a somewhat wider dark stripe and then several narrower and very distinct dark stripes which grow lighter ventrally. Distribution. Coastal Roach are restricted to coastal watersheds, largely west of the coastal mountain range. The northernmost native population is in the Navarro River, although introduced Coastal Roach exist in the Eel River. Excluding the Gualala River (see below), watersheds south of the Navarro River do not support any Roach populations until the Russian River. Populations are then found intermittently from the Russian River south to the Tomales Bay region. Scattered Coastal Roach are found in streams around San Francisco Bay but are once again absent from coastal watersheds to the south until the Salinas and Pajaro river watersheds of the Monterey region. The population in Soquel Creek (Santa Cruz Co.) is probably introduced from the Salinas River, while the origin of the population in the Cuyuma River (San Luis Obispo and Santa Barbara counties) is uncertain but may be native (Moyle 2002). Coastal Roach are widespread in the Eel River, from an introduction in the early1970s. This introduction was noted by Moyle (2002) and Jones (2002) and is now confirmed by Baumsteiger et al. (2017) and A. Kinziger (pers. comm. 2017). Studies show that Eel River fish originated from the Russian River mainstem and not the recently connected East Fork (via the Potter Valley Project - Power et al. 2015). Status. Due to its wide distribution throughout northern California coastal drainages, the species appears to be in little danger of extinction. However, subspecies or distinct population segments (see below) may be at risk within their particular watershed(s)., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 230-231, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Murphy, G. (1948) Distribution of Fariation of the Roach (Hesperoleucus) in the Coastal Region of California. M. A. Thesis, University of California, Berkeley. 55 pp.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Jones, W. J., Quelvog, B. D. & Bernardi, G. (2002) Morphological and genetic analysis of the Red Hills roach (Cyprinidae: Lavinia symmetricus). Conservation Genetics, 3, 261 - 276.","Power, M. E., Bouma-Gregson, K., Higgins, P. & Carlson, S. M. (2015) The thirsty Eel: summer and winter flow thresholds that tilt the Eel River of northwestern California from salmon-supporting to cyanobacterially degraded states. Copei a, 2015, 200 - 211."]}

Hesperoleucus venustus subditus Snyder, 1913 Southern Coastal Roach Holotype: USNM 74475. Originally described as Hesperoleucus subditus from Pajaro Basin, San Benito and Santa Clara Cos., CA. Paratypes: BMNH 1900.9.29.176; CAS-SU 22489; USNM 49579, 75338���40, 75342���43, 75345��� 50. Unlike previous listings by Snyder (1913), Tomales Bay (H. venustus) and Monterey Bay (H. subditus) species are genomically part of a southern subspecies of Coastal Roach (Baumsteiger et al. 2017). Previous works have recognized these individuals as distinct lineages both morphologically and genetically (Murphy 1948; Hopkirk 1973; Aguilar & Jones 2009). So while their distinctiveness seems consistent, their taxonomic level has not been properly defined till now. Description. Adult total length is typically 55���100 mm, up to 125 mm. Roach typically have 7���10 dorsal rays (average 9), 6���8 anal rays (average 7), and 47���62 lateral line scales (average 54) (Hopkirk 1973). The remaining characters vary between different DPSs proposed within this subspecies (see below), including thickness of caudal peduncle, fin length, body size, and snout shape (Snyder 1913; Murphy 1948; Hopkirk 1973). Distribution. Southern Coastal Roach are restricted to the drainages of Tomales Bay/northern SF Bay in the north and Monterey Bay in the south. There are no records of Roach being present in watersheds between these two systems but thorough sampling is needed to be sure of their absence. Status. Although both DPSs within this subspecies are considered to be Near-threatened (see below), the subspecies as a whole seems fairly abundant in their native ranges (Moyle et al. 2015). However, this is still probably a small subset of the original range of this subspecies, given the current geographic divide between the two DPSs within the subspecies. Both are found in areas where freshwater sources are limited and human use is high, making them candidates for decline through water manipulation. Additionally, most locations with this subspecies are relatively small, isolated creeks, making climate change an increasing problem for their persistence. Tomales Roach. Snyder (1913) described Hesperoleucus venustus from the Russian River, Tomales Bay tributaries, and San Francisco Bay tributaries. Hopkirk (1973) recognized that Roach from these three regions were different from each other but similar enough to California Roach that he declared H. venustus to be a synonym of H. s. symmetricus. He did suggest, however, that individuals in Tomales Bay tributaries were distinct enough to be recognized as a subspecies. The genetic studies of Jones (2001), Aguilar & Jones (2009) and Baumsteiger et al. (2017) indicated that populations from Tomales Bay tributaries are distinct, which we recognize here. Jones (2001) found populations of Tomales Roach from Lagunitas and Walker creeks shared nuclear DNA allele frequencies but were reciprocally monophyletic for mitochondrial DNA haplotypes. Aguilar & Jones (2009) determined from the mtDNA analysis of fish from Lagunitas and Walker Creeks that they were clearly a distinct lineage from CA Roach. Microsatellites, however, were not as definitive, with one analysis finding weak support for Tomales Roach as a distinct lineage, while another analysis found that, although distinguishable, individuals from Lagunitas and Walker Creeks clustered with Monterey Roach. Our genomic analysis (Baumsteiger et al. 2017) provided further clarity, showing that Tomales Bay locations are clearly definable within the Southern Coastal Roach subspecies, as a distinct population segment. Description. Tomales Roach look similar to other members of the CA Roach complex but are most similar to Russian River Roach (Table 1). Adult total length is typically 55���100 mm, up to 125 mm. Like Russian River Roach, Tomales Roach differ from CA Roach in having a more trim, slender body, a more pointed snout, a more slender caudal peduncle, and longer fins. Tomales Roach have 8���10 dorsal rays (average 9), 6���7 anal rays (average 7), and 47���62 lateral line scales (average 54) (Hopkirk 1973). In tributaries to San Francisco Bay, Snyder (1905, 1908) demonstrated that sex could be established by using the ratio of pectoral fin length to body length. Males exhibited a ratio of> 0.21 while females bore pectoral fins between 0.16 and 0.20 the length of their body. Both sexes exhibit bright orange and red breeding coloration on the operculum, chin and the base of the paired fins. Males may also develop numerous small breeding tubercles (pearl organs) on the head (Murphy 1943). Distribution. Tomales Roach are restricted to the Lagunitas Creek and Walker Creek drainages of western Marin County. Roach of uncertain taxonomic affinity have also been reported from Pine Gulch Creek, tributary to Bolinas Lagoon (Murphy 1948) and Salmon Creek, Marin Co. (Moyle 2002). However, a 1997 survey of Pine Gulch Creek recorded no Roach present (Fong 1999). Status. There is no indication that Tomales Roach in Walker and Lagunitas Creeks are in decline, but no regular surveys exist. Dams and diversions, however, have altered flows and habitat. They are currently listed as a state Species of Special Concern (IUCN Status, Near-threatened) but do not appear to be in immediate danger of extinction as a DPS. Fragmentation of populations, along with habitat alterations, may be limiting distribution and abundance (Moyle et al. 2015). Increasing urban water demands, coupled with predicted climate change impacts, are likely to lead to more widespread drying of stream segments and elimination of Roach populations. As a result, Moyle et al. (2013) considered Tomales Roach as ���highly vulnerable��� to extinction by 2100 from the effects of climate change. Moyle et al. (2015) considered it to be a species of ���moderate concern���, not facing immediate extinction threats. Monterey Roach. The Monterey Roach has been of interest because of its frequent hybridization with Monterey Hitch (L. e. harengus) (Miller 1945; Avise et al. 1975) but it retains its identity as a distinct taxon (Baumsteiger et al. 2017). Its isolation from other populations within the subspecies is of note, indicating that should be managed as a distinct population segment. Description. Monterey Roach are similar in appearance to CA Roach (see description) but can be distinguished by having fewer dorsal (7���9; average 8) and anal fins rays (6���8; average 7), fewer scales in the lateral line, slightly shorter fins, a slightly more robust body and a thicker caudal peduncle (Snyder 1913; Murphy 1948; Hopkirk 1973). Distribution. Monterey Roach are widely distributed in the Pajaro, Salinas, and San Lorenzo River systems as well as Soquel Creek, all tributaries to Monterey Bay on the central coast of California. Pescadaro Creek, just north of Monterey Bay, also contains Monterey Roach. Within the Pajaro watershed, Monterey Roach no longer occur in the mainstem but are present in Uvas Creek, Llagas Creek upstream of Chesbro Reservoir, the North Fork of Pacheco Creek upstream of Pacheco Reservoir, Arroyo Dos Picachos and in the San Benito River and its tributaries, including Tres Pinos, Laguna, and Clear Creeks (Smith 2007). In the Salinas River system, Roach also have been extirpated from the mainstem and now occur primarily in tributaries such as Arroyo Seco (J.J. Smith, pers. comm. 2009) and Gabilan Creek, although recent survey information is lacking. Status. Monterey Roach are apparently still numerous and widely distributed in smaller streams in much of their native range, but the DPS has been largely extirpated from the mainstem Pajaro and Salinas River systems due to habitat alteration, degraded water quality and quantity and alien species (Smith 1982, 2007). Long-term trends are not known but populations are likely fewer and more fragmented than historically. Moyle et al. (2015) listed them as a state Species of Special Concern (IUCN status of Near-threatened) and noted they were ���highly vulnerable��� to climate change due to potential reductions in stream flow throughout the watersheds (Moyle et al. 2013)., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 233-234, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Murphy, G. (1948) Distribution of Fariation of the Roach (Hesperoleucus) in the Coastal Region of California. M. A. Thesis, University of California, Berkeley. 55 pp.","Hopkirk, J. D. (1973) Endemism in fishes of the Clear Lake region of central California. University of California Publications in Zoology, 96, 1 - 160.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern","Jones, W. J. (2001) DNA Sequence Divergence and Speciation in Two California Minnows (Cyprinidae: Lavinia exilicauda and L. symmetricus). PhD Dissertation, University of California, Santa Cruz. Ca. 150 pp.","Snyder, J. O. (1905) Notes on the fishes of the streams flowing into the San Francisco Bay. Report for U. S. Bureau Fisheries for 1904, 327 - 338.","Snyder, J. O. (1908) The fishes of the coastal streams of Oregon and northern California. Bulletin, U. S. Bureau of Fisheries, 27, 153 - 189.","Murphy, G. (1943) Sexual dimorphism in the minnows Hesperoleucus and Rhinichthys. Copeia, 1943, 187 - 188.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Fong, D. (1999) 1997 California Freshwater Shrimp (Syncaris pacifica) Surveys within Point Reyes National Seashore and Golden Gate National Recreation Area (GGNRA) Prepared for the GGNRA, Division of Resource Management and Planning, and the U. S. Fish and Wildlife Service, Endangered Species Permits, Ecological Services, Portland Regional Office. 30 pp.","Moyle, P. B., Kiernan, J. D., Crain, P. K. & Quinones, R. M. (2013) Climate change vulnerability of native and alien freshwater fishes of California: a systematic assessment approach. PLoS One, 8 (5), e 63883.","Miller, R. R. (1945) The status of Lavinia ardesiaca, a cyprinid fish from the Pajaro-Salinas River basin, California. Copeia, 1945, 197 - 204.","Avise, J. C., Smith, J. J. & Ayala, F. J. (1975) Adaptive differentiation with little genic change between two native California minnows. Evolution, 29, 411 - 426.","Smith, J. J. (2007) Steelhead Distribution and Ecology in the Upper Pajaro River System and Mainstem Pajaro River. Unpublished Report 7, Department of Biology, San Jose State University. Ca. 40 pp.","Smith, J. J. (1982) Fishes of the Pajaro River system. University of California Publications in Zoology, 115, 83 - 170."]}

Hesperoleucus parvipinnis Snyder, 1913 Gualala Roach Holotype: USNM 74466. Originally described from Gualala River, Sonoma Co., CA. Paratypes: CAS-SU 14903. Gualala Roach were first collected by Snyder (1908) who recognized them as Rutilus symmetricus but noted they bore ���a distinctive local stamp by which they can be recognized without difficulty (p 175).��� He later (1913) described them as H. parvipinnis. Murphy (1948) suggested these individuals constituted a subspecies not a species. Aguilar & Jones (2009) found them to be strongly genetically different from any other Roach along the California coast, suggesting they were more than a subspecies. Baumsteiger et al. (2017) found individuals within the Gualala system to be distinctive at the species level in all analyses, with pairwise F ST estimates between these individuals and Northern Roach to be the highest in the study (0.895). Thus the consensus reconfirms what Snyder (1913) proposed, that Gualala Roach are indeed a distinct species within Hesperoleucus. Description. Adult Gualala Roach are small (typically 50���80 mm) and bronzy in color, similar in general appearance to California Roach (H. symmetricus). However, they differ from other Roach species by having smaller scales (54���65 along the lateral line), shorter, more rounded fins, a shorter snout (in relation to head length) and a more robust body (Table 1). Gualala Roach have 7���8 dorsal rays (average 8) and 6���8 (average 7) anal rays (Hopkirk 1973). Snyder (1913 p. 66) described Gualala Roach as having ���a light lateral stripe two scales wide extending from upper edge of gill opening to base of caudal and entirely above the lateral line; below is a somewhat wider dark stripe, which in turn is followed by several narrower and very distinct dark stripes which grow lighter ventrally.��� Distribution. Gualala Roach are confined to the Gualala River and its tributaries in Sonoma Co. They are the most abundant fish species in the South and Wheatfield forks and in most headwater streams (Entrix 1992; DeHaven 2008). Occurrence is greatly reduced in the colder North Fork (Parker and Pool 1964; CDFG 1991) and in the mainstem below its confluence with the North Fork (Kimsey 1953; DeHaven 2008). Status. Historically, Gualala Roach were present throughout the Gualala River basin (Higgins 1997). Agency salmonid surveys indicate Roach may have increased in abundance due to habitat alterations favorable to warm water-tolerant species (Higgins 1997). Moyle et al. (2013) rate this species as ���highly vulnerable��� to extinction due to climate change. Statewide this is a species of Special Concern (IUCN status, Near-threatened) because of its limited distribution and increasing threats from vineyard and rural residential development, including introductions of non-native fishes (Moyle et al. 2015)., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on page 236, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Snyder, J. O. (1908) The fishes of the coastal streams of Oregon and northern California. Bulletin, U. S. Bureau of Fisheries, 27, 153 - 189.","Murphy, G. (1948) Distribution of Fariation of the Roach (Hesperoleucus) in the Coastal Region of California. M. A. Thesis, University of California, Berkeley. 55 pp.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Hopkirk, J. D. (1973) Endemism in fishes of the Clear Lake region of central California. University of California Publications in Zoology, 96, 1 - 160.","Entrix Inc. (1992) South Fork Gualala River Fishery Resource Stud y. Prepared for the Sea Ranch Water Company. Walnut Creek, CA. 63 pp.","DeHaven, R. W. (2008) Adult and Juvenile Steelhead Population Surveys, Gualala River, California, 2008. Unpublished report, prepared by the author December 31, 2008, for use by agencies, groups and individuals involved in steelhead recovery efforts. 47 pp.","Parker, C. & Pool, R. (1964) North Fork Gualala River Stream Survey, North Fork Gualala River surveyed by foot and automobile from the mouth to the headwaters, a total of 15 miles. September 17 - 18, 1964. File report, California Department of Fish and Game, Region 3, Yountville, CA. 2 pp.","Kimsey, J. B. (1953) Population Sampling of Three North Coastal Streams Closed to Summer Trout Fishing - 1952 Season. First Progress Report. California Department of Fish and Game, Sacramento. 15 pp.","Higgins, P. (1997) Gualala River Watershed Literature Search and Assimilation. Redwood Coast Land Conservancy, Gualala CA. 57 pp.","Moyle, P. B., Kiernan, J. D., Crain, P. K. & Quinones, R. M. (2013) Climate change vulnerability of native and alien freshwater fishes of California: a systematic assessment approach. PLoS One, 8 (5), e 63883.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern"]}

Hesperoleucus symmetricus symmetricus (Baird & Girard, 1854) Central California Roach This subspecies is consistent with the descriptions and distribution of the species itself, minus the Red Hills region. According to Baumsteiger et al. (2017), population structure is available within the subspecies, but only one lineage (see below) stood out in the genomic analyses and therefore would rise to the level of a DPS by our estimation. However, the presence of population structure and morphological differences (Brown et al. 1992) within the subspecies indicates isolation/adaptation among locations that, if fully investigated, may warrant additional DPS designations. We would argue this should be investigated on a case-by-case basis to evaluate the level of isolation and potential threat to each population so as to not overuse the DPS designation within the U.S. Endangered Species Act. Kaweah Roach. The current population of Kaweah Roach is highly isolated in the Kaweah River system, at the southern end of the CA Roach range. Some morphometric characters are distinctive in Kaweah Roach, including a high prevalence of chisel-lip (Brown et al. 1992). Although found to be genetically (Aguilar & Jones 2009) and genomically distinct (Baumsteiger et al. 2017), these determinations were only discovered once analyses were restricted to CA Roach samples. Therefore, we believe Kaweah Roach is a unique evolutionary lineage within CA Roach, representing a distinct population segment (DPS) that should be managed accordingly. Description. Kaweah Roach are largely similar to the description for CA Roach, with slightly different morphometric and meristic mean counts reported (Brown et al. 1992). This includes a higher prevalence of chisellip, similar to Red Hills Roach. Distribution. This distinct population segment appears to be isolated in the Kaweah River above Kaweah Reservoir. Historically it may have occurred on the Valley floor, in places such as Tulare Lake or in the lower reaches of nearby river systems such as the Kings River. Such access is now denied by dams, diversions, and altered habitats. Status. This DPS is reasonably safe, at least in the short run (50 years). The upper half of the range within the Kaweah River is bounded by Kings Canyon National Park and as such receives special protection. The lower half, which flows mostly through private lands before ending up in Kaweah Reservoir, is less secure. Individuals here may be at risk from upstream movement of invasive species such as Largemouth Bass, Micropterus salmoides, a factor which will intensify with climate change and warmer temperatures.

Hesperoleucus venustus navarroensis Snyder, 1913 Northern Coastal Roach Holotype: USNM 74477. Originally described as Hesperoleucus navarroensis from Navarro River, near Philo, Mendocino Co., CA. Paratype: CAS-SU 22488. Snyder (1913) described two species as occurring within the range of this subspecies: H. navarroensis from the Navarro River basin, and H. venustus, from a mixture of Russian River, Tomales Bay, and San Francisco Bay regions. Hopkirk (1973) disagreed with Snyder, placing Russian River individuals within the subspecies H. s. navarroensis and downgrading Tomales and San Francisco Bay individuals to population status. Aguilar & Jones (2009) found genetic distinctiveness for a Navarro/Russian grouping and a Tomales/SF Bay grouping within their microsatellite analyses. They also found some mtDNA distinctiveness for each location proposed (Navarro, Russian, and Tomales/SF Bay). Baumsteiger et al. (2017) found clear genomic distinctions between Navarro/ Russian and Tomales/SF Bay individuals, consistent with other studies, supporting our assertion of two subspecies within the Coastal Roach species. Description. Like other Coastal Roach, members of this subspecies are relatively trim, with a slender caudal peduncle, short, pointed snout, and variable length fins (dependent on which region is sampled). Individuals are typically 80-120 mm total length, with a large, conical head. Eyes are large and the mouth is sub-terminal. The dorsal fin has 7���9 rays (average 8) and the anal fin has 6���9 rays (average 8). Most pharyngeal teeth (0,5���4,0) have curved tips which overhang slightly. Scales are small, numbering 47���63 (average 54) along the lateral line. Fish appear darker on the upper half while the lower half is much lighter. Northern Coastal Roach differ from Gualala Roach in having fewer anal rays (Hopkirk 1973) and one less row of scales above the lateral line (Snyder 1913). Distribution. Northern Coastal Roach are generally restricted to the Navarro River and Russian River basins. There are no records of Roach being found in watersheds between these basins. The exact distribution within the two watersheds is poorly documented but they appear to be present wherever water quality is suitable and alien predators are absent. The introduced population in the Eel River constitutes a major range expansion north. Status. Because this subspecies is restricted to just two watersheds in its native range, it is highly subject to localized effects of farming, ranching, water diversions, dams, and other perturbations. However there are no reports of dwindling numbers in these systems, suggesting that, for now, populations of this subspecies are relatively robust. Russian River Roach. The Russian River Roach was originally described as a population of H. venustus by Snyder (1913), along with populations in streams flowing intoTomales and San Francisco bays. Following the downgrading of this group to subspecies status, Russian River Roach were later lumped with Navarro Roach as H. s. navarroensis by Hopkirk (1973). This move was confirmed genetically by Aguilar & Jones (2009). The recent analysis by Baumsteiger et al. (2017) found them to be part of the Northern subspecies of Coastal Roach but as a distinct population segment (DPS). Hierarchical genomic analyses revealed Eel River individuals cluster (are synonymous) with Russian River individuals in all analyses. An analysis using only fish from the two watersheds showed population structure at each of three sampling locations within the Russian River whereas within the Eel River there was no population structure among the four locations sampled. This means that the Eel River population is derived from a very recent introduction from the Russian River. Description. Russian River Roach have a trim, slender body, a somewhat pointed snout, a slender caudal peduncle. The fins are somewhat longer than those of other Coastal Roach. They have 8���9 dorsal rays (average 9) and 7���9 anal rays (average 8); there are 8���11 gill rakers (average 10) and 47���60 scales in the lateral line (average 53) (Hopkirk 1973). Individuals rarely exceed 120 mm total length. Minor morphological differences were observed between Russian River and Navarro Roach by Snyder (1913) and Murphy (1948). Differences included a higher prevalence of individuals with 9 dorsal rays in the Russian rather than the 8 generally seen in Navarro (see Table 1). Distribution. Russian River Roach are restricted to the Russian River and its tributaries in Sonoma Co. as well as to the Eel River watershed. They are common in the middle sections of many tributary creeks, including Mark West, Santa Rosa, Maacama, Austin, and Big Sulphur and Pieta (Moyle et al. 2015). Individuals are also present in the main stem Russian River (Pintler & Johnson 1958; Cook 2005). They become increasingly rare in the lower sections of the main river, where their downstream limit appears to be Duncan���s Mill, just above the main estuary (Moyle et al. 2015). However, Goodwin et al. (1993) found that Roach seemed to move down into the estuary during the summer and return upstream in the fall. Within the Eel River watershed, fish are widespread and abundant, with Brown & Moyle (1997) finding individuals at 230 of 412 (56%) sampling locations in the watershed. Status. Little is known of Russian River Roach abundance trends in their native habitat because few recent, systematic surveys have been conducted in tributary streams where they are most abundant. In the Russian River, gradual loss of tributary populations through dewatering and other factors, combined with mainstem river changes that reduce connectivity among tributaries may limit distribution and reduce abundance. The DPS is currently listed as a state Species of Special Concern (IUCN, Near-threatened) because of a limited distribution, the presence of dams and diversions in the Russian River watershed, and a poorly understood status (Moyle et al. 2015). Eel River fish were not included in the designation. The increasingly stressful conditions in the Russian River and its tributaries led Moyle et al. (2013) to rate the Russian River Roach as ���highly vulnerable��� to extinction in its native range as the result habitat loss due to climate change, if present trends continue. Navarro Roach. Hopkirk (1973) was the first to suggest that the original species/subspecies described by Snyder (1913) should include Russian River individuals. Aguilar & Jones (2009) and Baumsteiger et al. (2017) later found genetic distinctiveness between individuals from the Navarro and Russian rivers, prompting their recognition as distinct population segments of Northern Coastal Roach. Description. Like other Coastal Roach, Navarro Roach are small, stout-bodied cyprinids with a narrow caudal peduncle, deeply forked tail, short snout, and small, rounded paired fins. They are typically less than 100 mm total length. The head is large and conical, the eyes large and the mouth is sub-terminal and slants downward. The dorsal fin is short (7���9 rays; average 8) and is positioned behind the insertion point of the pelvic fin. The anal fin has between 6���9 rays. Pharyngeal teeth (0,5���4,0) have curved tips which overhang grinding surfaces of moderate size. Scales are small, numbering 47���63 along the lateral line and 32���38 before the dorsal fin. Individuals are usually dark on the upper half of their bodies, ranging from a shadowy gray to a steel blue, while the lower half is much lighter, usually a dull white/silver color. Snyder (1913) described them as having a light lateral stripe approximately two scales wide extending from upper edge of the gill opening to the base of tail and entirely above the lateral line. Below this line is a somewhat wider dark stripe and then several narrower and very distinct dark stripes which grow lighter ventrally. As mentioned previously, small morphological differences are found between Russian and Navarro populations (Table 1). Distribution. Navarro Roach are confined to the Navarro River and its tributaries within Mendocino Co. (including Greenwood Creek) but are widely distributed within the watershed, from the mouth throughout the river and its tributaries (Feliciano 2004). Status. The Navarro Roach, through its original lumping with CA Roach, was a state Species of Special Concern with an IUCN status of Near-threatened (Moyle et al. 2015). However with its new taxonomic status, a reevaluation will eventually be necessary, despite being widespread and abundant within its limited range. Most of the Navarro River watershed was logged and converted to orchards and vineyards in the 20 th century. While removal of big trees created more of the warm-water habitat preferred by Navarro Roach, diversion of water has greatly reduced flows, presumably countering the effects of tree removal. Because of these factors, Moyle et al. (2013, 2015) considered it highly vulnerable to extinction in the next century from climate change and other factors., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 231-232, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Hopkirk, J. D. (1973) Endemism in fishes of the Clear Lake region of central California. University of California Publications in Zoology, 96, 1 - 160.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Murphy, G. (1948) Distribution of Fariation of the Roach (Hesperoleucus) in the Coastal Region of California. M. A. Thesis, University of California, Berkeley. 55 pp.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern","Pintler, H. E. & Johnson, W. C. (1958) Chemical control of rough fish in the Russian River drainage, California. California Fish and Game, 44, 91 - 124.","Cook, D. (2005) Chinook Salmon Spawning Study, Russian River, Fall 2002 - 2004. File Report. Sonoma County Water Agency, Santa Rosa, CA. 20 pp.","Goodwin, P., Cuffe, C. K., Nielsen, J. L. & Light, T. (1993) Russian River Estuary Study, 1992 - 1993. Department of Planning, Sonoma County, California, U. S. A. 186 pp.","Brown, L. R. & Moyle, P. B. (1997). Invading species in the Eel River, California: successes, failures, and relationships with resident species. Environmental Biology of Fishes, 49, 271 - 291.","Moyle, P. B., Kiernan, J. D., Crain, P. K. & Quinones, R. M. (2013) Climate change vulnerability of native and alien freshwater fishes of California: a systematic assessment approach. PLoS One, 8 (5), e 63883.","Feliciano, J. B. (2004) A Study of the Salmonid Populations in the Navarro River Watershed: Tracking Declines and Tools for Monitoring. Ph. D. Dissertation, University of California, Davis. 80 pp."]}

Hesperoleucus symmetricus symmetricus (Baird & Girard, 1854) Central California Roach This subspecies is consistent with the descriptions and distribution of the species itself, minus the Red Hills region. According to Baumsteiger et al. (2017), population structure is available within the subspecies, but only one lineage (see below) stood out in the genomic analyses and therefore would rise to the level of a DPS by our estimation. However, the presence of population structure and morphological differences (Brown et al. 1992) within the subspecies indicates isolation/adaptation among locations that, if fully investigated, may warrant additional DPS designations. We would argue this should be investigated on a case-by-case basis to evaluate the level of isolation and potential threat to each population so as to not overuse the DPS designation within the U.S. Endangered Species Act. Kaweah Roach. The current population of Kaweah Roach is highly isolated in the Kaweah River system, at the southern end of the CA Roach range. Some morphometric characters are distinctive in Kaweah Roach, including a high prevalence of chisel-lip (Brown et al. 1992). Although found to be genetically (Aguilar & Jones 2009) and genomically distinct (Baumsteiger et al. 2017), these determinations were only discovered once analyses were restricted to CA Roach samples. Therefore, we believe Kaweah Roach is a unique evolutionary lineage within CA Roach, representing a distinct population segment (DPS) that should be managed accordingly. Description. Kaweah Roach are largely similar to the description for CA Roach, with slightly different morphometric and meristic mean counts reported (Brown et al. 1992). This includes a higher prevalence of chisellip, similar to Red Hills Roach. Distribution. This distinct population segment appears to be isolated in the Kaweah River above Kaweah Reservoir. Historically it may have occurred on the Valley floor, in places such as Tulare Lake or in the lower reaches of nearby river systems such as the Kings River. Such access is now denied by dams, diversions, and altered habitats. Status. This DPS is reasonably safe, at least in the short run (50 years). The upper half of the range within the Kaweah River is bounded by Kings Canyon National Park and as such receives special protection. The lower half, which flows mostly through private lands before ending up in Kaweah Reservoir, is less secure. Individuals here may be at risk from upstream movement of invasive species such as Largemouth Bass, Micropterus salmoides, a factor which will intensify with climate change and warmer temperatures., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 228-229, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Baird, S. F. & Girard, C. F. (1854) Descriptions of new species of fishes collected in Texas, New Mexico and Sonora, by Mr. John H. Clark, on the US and Mexican boundary survey and in Texas by Capt. Stewart Van Vliet, Proceedings, Academy of Natural Sciences of Philadelphia, 7, 24 - 29.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Brown, L. R., Moyle, P. B., Bennett, W. A. & Quelvog, B. D. (1992) Implications of morphological variation among populations of California roach Lavinia symmetricus (Cyprinidae) for conservation policy. Biological Conservation, 62, 1 - 10.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381."]}

Hesperoleucus mitrulus Snyder, 1913 Northern Roach Holotype: USNM 74474. Originally described from Drew Creek, Lake Co., OR. Paratypes: CAS-SU 22491; UMMZ 139019. Northern Roach were first collected in 1898 by C. Rutter as Rutilus symmetricus (Rutter 1908). Rutter (1908: 139) stated “We have but few small specimens of this form, the longest being but 3 inches long. They were taken in North Fork Pitt (sic) River near Alturas and at the mouth of Joseph Creek, several hundred miles from where any other specimens of symmetricus have been taken. The form may prove to not to be symmetricus, but we cannot identify it otherwise with the material at hand.” In 1913, Snyder described the Goose Lake drainage fish as Hesperoleucus mitrulus. The first inclusion of Roach from the Pit River as part of H. symmetricus was by Hubbs et al. (1979). Genetically, Aguilar & Jones (2009) found that individuals from these locations were distinct from all other CA Roach. Our genomic study (Baumsteiger et al. 2017) also found all individuals collected from these locations to be distinct at the species-level in every analysis. Description. Northern Roach are small (adult size typically 50–100 mm TL) bronzy cyprinids, similar in appearance to the California Roach (Moyle 2002; Fig. 3). They have a robust body, deep caudal peduncle, short snout and short rounded fins. Northern Roach differ from CA Roach in having short rounded fins and “cup-like” scales (Snyder 1913; Table 1). Snyder (1908) examined 20 fish from Drew Creek, Oregon and all individuals had 8 dorsal rays and 7 anal rays. Snyder found that males had longer, larger fins than did females, especially pectoral fins; the sexes could be differentiated by the ratio of pectoral fin length to body length. Distribution. The original collections by Rutter (1908) consisted of the North Fork of the Pit River and the mouth of Joseph Creek. Snyder (1908) collected widely in the upper Pit River, Goose Lake basin of California and Oregon, and in the Summer, Abert, Harney and Warner basins of Oregon but only found Roach in tributaries to Goose Lake, Lake Co., Oregon. The first inclusion of individuals from the Pit River as part of this species was by Hubbs et al. (1979). While no mention is made of a range extension for the taxon, it is assumed this change was precipitated by the 1934 collection of 19 roach in the North Fork Pit River near Alturas, Modoc County (unpublished field notes and collections of Carl Hubbs at the Univ. of Michigan as reported in Reid et al. 2003). In the most comprehensive sampling of the Pit system in California to date, Moyle and Daniels (1982) found Roach at only 8% of 261 collection sites. Northern Roach were found in only three widely separated drainages: (1) Ash–Rush–Willow Creek drainage, Lassen/Modoc Co., (2) Bear Creek, tributary to the Fall River, Shasta Co. and (3) Beaver Creek, Lassen Co. All locations are above Pit River Falls (Shasta Co.) which divides the Pit River Basin into upper and lower drainages. The falls are a barrier to fish movement (Kinziger et al. 2016). Historically, the waterfall was the northern range limit for some Sacramento River fishes, such as Tule Perch, Hysterocarpus traskii (Moyle 2002). Roach below the falls had unimpeded access to the Sacramento River system and are assumed to be H. s. symmetricus. However, genetic and morphometric studies have not been conducted and relationships remain uncertain. Status. Northern Roach have been uncommon in California ever since they were first collected in 1898, although by that time much of the Pit River drainage had been heavily altered by grazing, agriculture and logging (Moyle & Daniels 1982). Subsequently, alien fish species that compete with or prey on Northern Roach (e.g. Green Sunfish, Lepomis cyanellus) became widely introduced. Their absence from some streams, however, may be the result of specific habitat requirements. Northern Roach prefer spring pools and swampy stream reaches, unlike the intermittent stream habitats where other species of Roach are found (S. Reid, pers. comm.). Thus in Lassen/Modoc Co., Roach are found in small numbers, inhabiting the weedy margins of streams and, in one case, an isolated spring pond (Moyle & Daniels 1982; S. Reid, pers. comm.). However a fairly recent survey of fishes in the Oregon portion of the Goose Lake watershed found Northern Roach to be relatively abundant in six small creeks (Scheerer et al. 2010). Moyle et al. (2015) rated the Northern Roach as a Species of Special Concern in California and the International Union for the Conservation of Nature (IUCN) lists their status as High Concern. Additionally, these Roach are highly vulnerable to extinction in the next century from climate change effects on their small streams (Moyle et al. 2013)., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 234-235, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin: with a study of their distribution and variation. Bulletin, U. S. Fish Commission, 27, 103 - 152.","Hubbs, C. L. Follett, W. I. & Dempster, L. J. (1979) List of the fishes of California. Occasional Papers of the California Academy of Sciences, 133, 1 - 51","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Snyder, J. O. (1908) The fishes of the coastal streams of Oregon and northern California. Bulletin, U. S. Bureau of Fisheries, 27, 153 - 189.","Reid, S. B., Cowan, W. & Byers, S. (2003) Mainstem upper Pit River fish surveys including the North and South Fork Pit Rivers, September 2002. File Report, U. S. Fish and Wildlife Service, Klamath Falls, Oregon. 53 pp","Moyle, P. B. & Daniels, R. A. (1982) Fishes of the Pit River system, and Surprise Valley region. University of California Publications in Zoology, 115, 1 - 82.","Kinziger, A. P., Hellmair, M., Fong, S. R., Goodman, D. H. & Kelsey, H. (2016) Evolution of rough sculpin (Cottus asperrimus) genetic divergence and late Quaternary displacement on the Hat Creek fault, California, USA. Conservation Genetics, 17, 1257 - 1267.","Scheerer, P. D., Gunckel, S. L., Heck M. P. & Jacobs, S. E. (2010) Status and distribution of native fishes in the Goose Lake Basin. Northwestern Naturalist, 91, 271 - 287.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern","Moyle, P. B., Kiernan, J. D., Crain, P. K. & Quinones, R. M. (2013) Climate change vulnerability of native and alien freshwater fishes of California: a systematic assessment approach. PLoS One, 8 (5), e 63883."]}

Lavinia and Hesperoleucus One conundrum within the Hitch–Roach species complex is uncertainty regarding whether species should be placed in one genus or two genera. While the complex was originally divided between Lavinia and Hesperoleucus, multiple examples of hybridization between the genera suggest they are more closely related than separate genera designations would normally indicate. For example, CA Roach are known to hybridize with Hitch in the Pajaro and Salinas rivers in Monterey Bay drainages (Miller 1945), in Coyote, Alameda, and Walnut Creeks in the San Francisco Bay region (Miller 1963; Leidy 1984, 2007, pers. comm.), and in Sacramento-San Joaquin drainages (Avise et al. 1975; Jones 2001). Hybrids were fertile in the Pajaro River; Avise et al. (1975) found 8% of Hitch examined to be F 1 hybrids and 5% to be backcrossed. Since it is unusual for two genera to be interfertile and because other endemic species in California do not appear to exhibit this behavior (Avise et al. 1975; Avise & Ayala 1976; Moyle & Massingill 1981; Moyle 2002), it has been proposed these two species belonged in a single genus. Aguilar & Jones (2009) found the two genera are closely related mtDNA lineages that could be considered to be one genus. Schönhuth et al. (2012), however, noted that while their mtDNA analyses supported this close relationship, their limited nuclear DNA results did not. Our recent study (Baumsteiger et al. 2017) found the strongest genomic signal coincided with splitting samples into two genera. We discovered, however, that samples from the Pit River (Northern Roach) were a confounding factor, having mixed historical ancestry with both genera. It was estimated in one analysis that approximately 80% of the Northern Roach genome clustered with Hitch whereas only 20% clustered with CA Roach. But all subsequent analyses found Northern Roach to be a distinct species, on par with currently identified species. Thus Northern Roach appears to be the result of an ancient hybrid speciation event (Mallet 2007) between ancestral Hitch and CA Roach but it is currently genomically different enough to warrant species status. It is our conclusion that despite hybridization between individuals in the different genera, the distinctiveness of each genus is still strongly maintained and prevalent throughout the genomes interrogated. If results were more admixed, the argument for a single genus would be more plausible. We recognize admixture between genera is uncommon but note that not all mutations lead to post-zygotic barriers (Abbott et al. 2013). Additionally, hybridization has been proposed between Roach/Hitch and other members of the subfamily Leuciscinae including Speckled Dace (Rhinichthys osculus), Arroyo Chub (Gila orcuttii), and the now extinct Thicktail Chub (Gila crassicauda) (Greenfield & Deckert 1973; Hopkirk 1973). Thus the frequency of hybridization suggests reproductive barriers are poorly developed in the endemic minnows of California. Secondly, although Northern Roach represent an ancient hybridization event and cluster primarily with individuals from the Lavinia genus (~80% but only in the admixture analysis), we cannot definitively say they are Lavinia. Northern Roach still strongly resemble CA Roach in appearance (see below), but this may simply be the result of natural selection for the CA Roach morphotype. Future phylogenetic analyses between all genera within the family may provide the solution but it remains, as yet, unavailable. Thus, without universal agreement between genomic analyses or improved criteria for taxonomically delineating hybrids, we default to the original genus assignment by Snyder (1913). Therefore, the species of Hesperoleucus are as described below.

Hesperoleucus symmetricus (Baird & Girard, 1854) California Roach Lectotype: USNM 191, present designation. This species was originally described as Pogonichthys symmetricus by Baird & Girard (1854) from specimens collected in the San Joaquin River at Fort Miller, Fresno Co., CA. Paralectotypes: USNM 440410; ANSP 5330–33; MCZ 1961; MNHN 0000–0353. The benchmark California Roach is based on Baird & Girard (1854). Individuals primarily occupy streams/ rivers flowing into California’s Central Valley, most of which are isolated to some degree. Groups of California Roach appear to become easily isolated from one another and adapted to local conditions, particularly following anthropomorphic changes to the landscape (Brown et al. 1992). Many of these isolated groups are distinguishable both morphologically (Brown et al. 1992) and genetically (Aguilar & Jones 2009; Baumsteiger et al. 2017). In fact, Baumsteiger et al. found every sample location of CA Roach within our study was distinctive, including support for a new subspecies and a distinct population segment (see below). Baumsteiger et al. (2017) also found California Roach from the Central Valley to be genomically distinct from those along the coast. All analyses show a clear break between these two species. Our conclusion is CA Roach sensu stricto should be limited to individuals from Central Valley drainages and the following formal description reflects this new delineation. Description. CA Roach (H. symmetricus) are small, stout-bodied cyprinids with narrow caudal peduncles and deeply forked tails. Adult individuals usually have total lengths less than 100 mm, although fish up to 150 mm TL have been observed (Moyle 2002). The head is large and conical, with large eyes and a mouth that is slightly subterminal and slants downward. Individuals in some locations, especially those in streams of the Sierra Nevada, develop a keratinous plate on the lower jaw, referred to as a “chisel lip”. The dorsal fin is short (8–10 rays) and is positioned behind the insertion point of the pelvic fin while the anal fin is also short, with 7–9 rays (Table 1). Pharyngeal teeth (0,5,4,0) have curved tips, which overhang grinding surfaces of moderate size. CA Roach are dark on the upper half of their bodies, ranging from a shadowy gray to a steel blue, while the lower half of the body is much lighter, usually a dull white/silver color. Scales are small, numbering 47–62 along the lateral line. Individuals tend to exhibit general (non-nuptial) sexual dimorphism (Snyder 1908; Murphy 1943). California Roach are highly variable in most of their characteristics across their range. Brown et al. (1992) measured 14 morphometric variables from eight watersheds around the San Joaquin Valley, including the southern end of the range (Table 1). Using these variables, a discriminant analysis correctly assigned 76% of the individuals to the watershed from which they were collected, suggesting locations were sufficiently isolated from one another to have local adaptations or phenotypic divergence. ……continued on the next page Distribution. Individuals were historically found in most tributaries to the Sacramento and San Joaquin Rivers, including tributaries in the extreme southern San Joaquin Valley (e.g. Kern River), as well as in fringe habitats (backwaters with dense riparian cover) along the main rivers (Moyle 2002; Fig. 1). Fish were rarely found in streams above 1000 m in elevation, presumably restricted by high gradients and natural barriers. In the Tuolumne River, the upstream limit was historically (and still is) Preston Falls at 885 m elevation. The historic distribution in the upper Sacramento River is poorly understood. Today CA Roach are absent from many streams as the result of habitat change and invasions of alien predators. For example, Moyle & Nichols (1973) and Brown & Moyle (1993) found they were absent from the Fresno River (Fresno Co.) and other tributaries to the San Joaquin River. They were also once common in the mainstem San Joaquin River but are absent today. Moyle (2002) documented their consistent presence in the river at Friant (Fresno Co.) until the construction of Friant Dam in the 1940s. CA Roach are absent from most of their historic range in the Cosumnes River watershed due to the invasion of piscivorous Redeye Bass, Micropterus coosae (Moyle et al. 2003). In the upper Yuba River watershed, they are largely absent as the result of 19 th century hydraulic mining, with the exception of one small tributary to the South Fork Yuba, Kentucky Ravine (Gard & Randall 2004). The range of CA Roach has also been expanded, presumably through ‘bait bucket’ introductions by anglers. For example, Hetch-Hetchy Reservoir (1,162 m) on the upper Tuolumne River, well above a series of natural barriers, supports a large pelagic population (P. Moyle, unpublished observations). This is unusual because CA Roach do not generally live in reservoirs, especially those with predatory fishes. Interestingly, individuals from this location were found to be admixed with Hitch (Baumsteiger et al. 2017), a factor which may explain their persistence. Soquel Creek and the Cuyama River in southern California support presumed introduced populations, although their origins are also uncertain (Moyle 2002). It is possible that the ancestor of the widely-distributed CA Roach gave rise to most other Roach species and subspecies but this has not yet been investigated phylogenetically. The ability of these fish to persist in small, often intermittent tributaries presumably led to their colonization of adjacent drainages in a number of areas through capture of interior headwater streams by erosion or tectonic movements (Snyder 1908, 1913; Murphy 1948; Moyle 2002). Because they are relatively intolerant of saline waters, dispersal to coastal streams could not have occurred through ocean waters. However, connections at low elevations may have been possible in some cases when sea levels were lower and estuaries joined the mouths of rivers (Moyle 2002). Similarly in the giant San Francisco Estuary, exchange between salt-water intolerant populations likely occurred during flood years when river outflows were high enough to create freshwater lenses in the surface waters of the estuary (Ayres 1854; Snyder 1905; Murphy 1948; Leidy 2007). During high water periods, fish may also have been able to disperse through flooded marshes on the fringes of the estuary. Today such opportunities for dispersal are largely absent (Moyle et al. 2012). Status. Moyle et al. (2015) list California Roach as a Species of Special Concern, with an IUCN status of Near–threatened. If CA Roach were a single interbreeding taxon, there would be little danger of immediate extinction because they occur in many streams over a wide area. However, the discovery that CA Roach comprise multiple lineages, some of which are under immediate threats, is of conservation concern. After all, the small, isolated populations that are the most likely to be extirpated also tend to be the most distinctive (Brown et al. 1992). Emerging appreciation for the variation within the taxon (Moyle et al. 1989; Brown et al. 1992; Jones 2001; Jones et al. 2002; Aguilar & Jones 2009; Baumsteiger et al. 2017) has highlighted the need to preserve populations endemic to specific watersheds. On a broader scale, Moyle et al. (2013) regarded the entire taxon as highly vulnerable to extinction in the next century due to massive changes to smaller streams brought on by climate change, the continuing effects of anthropogenic land and water use, and the spread of invasive species., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 224-228, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Baird, S. F. & Girard, C. F. (1854) Descriptions of new species of fishes collected in Texas, New Mexico and Sonora, by Mr. John H. Clark, on the US and Mexican boundary survey and in Texas by Capt. Stewart Van Vliet, Proceedings, Academy of Natural Sciences of Philadelphia, 7, 24 - 29.","Brown, L. R., Moyle, P. B., Bennett, W. A. & Quelvog, B. D. (1992) Implications of morphological variation among populations of California roach Lavinia symmetricus (Cyprinidae) for conservation policy. Biological Conservation, 62, 1 - 10.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Snyder, J. O. (1908) The fishes of the coastal streams of Oregon and northern California. Bulletin, U. S. Bureau of Fisheries, 27, 153 - 189.","Murphy, G. (1943) Sexual dimorphism in the minnows Hesperoleucus and Rhinichthys. Copeia, 1943, 187 - 188.","Moyle, P. B. & Nichols, R. D. (1973) Ecology of some native and introduced fishes of the Sierra Nevada foothills in central California. Copeia, 1973, 478 - 490.","Brown, L. R. & Moyle, P. B. (1993) Distribution, ecology, and status of the fishes of the San Joaquin River drainage, California. California Fish and Game, 79, 96 - 114.","Moyle, P. B., Crain, P. K., Whitener, K. & Mount, J. F. (2003) Alien fishes in natural streams: fish distribution, assemblage structure, and conservation in the Cosumnes River, California, USA. Environmental Biology of Fishes, 68, 143 - 162.","Gard, M. & Randall, P. (2004) Setting priorities for native fish conservation: An example from the South Yuba River watershed. California Fish and Game, 90, 1 - 12.","Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72.","Murphy, G. (1948) Distribution of Fariation of the Roach (Hesperoleucus) in the Coastal Region of California. M. A. Thesis, University of California, Berkeley. 55 pp.","Ayres, W. O. (1854) Descriptions of new species of cyprinoids. Originally published in Daily Placer Times and Transcript, May 1854 and reprinted in 1855 in Proceedings of California Academy of Sciences, 1, 18 - 22.","Snyder, J. O. (1905) Notes on the fishes of the streams flowing into the San Francisco Bay. Report for U. S. Bureau Fisheries for 1904, 327 - 338.","Leidy, R. A. (2007) Ecology, Assemblage Structure, Distribution, and Status of Fishes in Streams Tributary to the San Francisco Estuary, California. San Francisco Estuary Institute, San Francisco. 194 pp.","Moyle, P. B., Hobbs, J. & O'Rear, T. (2012) Fishes. In A. Palaima, ed. Ecology, Conservation and Restoration of Tidal Marshes: The San Francisco Estuary. University of California Press. Berkeley. pp. 161 - 173.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern","Moyle, P. B., Williams, J. E. & Wikramanayake, E. D (1989) Fish Species of Special Concern of California. California Department of Fish and Game. Sacramento. 272 pp.","Jones, W. J. (2001) DNA Sequence Divergence and Speciation in Two California Minnows (Cyprinidae: Lavinia exilicauda and L. symmetricus). PhD Dissertation, University of California, Santa Cruz. Ca. 150 pp.","Jones, W. J., Quelvog, B. D. & Bernardi, G. (2002) Morphological and genetic analysis of the Red Hills roach (Cyprinidae: Lavinia symmetricus). Conservation Genetics, 3, 261 - 276.","Moyle, P. B., Kiernan, J. D., Crain, P. K. & Quinones, R. M. (2013) Climate change vulnerability of native and alien freshwater fishes of California: a systematic assessment approach. PLoS One, 8 (5), e 63883."]}

Hesperoleucus symmetricus serpentinus Baumsteiger and Moyle, new subspecies Red Hills Roach (Fig. 2) Holotype: Museum of Wildlife and Fish Biology, University of California, Davis, WFC3050, 51.8 mm SL, Horton Creek, Tuolumne County, CA, July 16, 2010. Paratypes. WFC3241���3244, 30.4���40.4 mm SL, same location and date as the holotype, The Red Hills Roach has strong support for a subspecies designation (e.g. Brown et al. 1992; Jones et al. 2002; Moyle 2002), especially given its increased isolation from other populations by New Don Pedro Reservoir (constructed in 1971), unique environmental constraints, and the prevalence (~97%) of the chisel-lip morphotype. Genetic and genomic analyses (Aguilar & Jones 2009; Baumsteiger et al. 2017) also found Red Hills Roach to be highly distinct, easily distinguishable from other CA Roach. In fact, our genomic study indicated a clear break between Red Hills Roach and all other CA Roach sampled in every analysis. With so many lines of support, subspecies status is clearly warranted. Diagnosis. Distinguished from other subspecies of H. symmetricus by their flattened body profile, smaller interorbital distance and fewer pectoral and pelvic rays (Jones et al. 2002; Table 1). The holotype has 8 dorsal rays, 8 anal rays, 12 pectoral rays, and 8 pelvic rays. Description. The Red Hills Roach is a genetically and morphologically distinct minnow adapted for living in small streams flowing through serpentine outcrops. They are typically small, less than 60 mm SL (Brown et al. 1992; Jones et al. 2002) and resemble CA Roach in their general appearance. Red Hills Roach exhibit an elevated frequency of ���chisel-lip���, a keratinous projection on the lower lip used for scraping algae (Brown et al. 1992). However there is substantial temporal variation in frequency of the chisel-lip condition amongst CA Roach (Jones et al. 2002). Spawning coloration appears to be similar to that of CA Roach but is more intense. W. J. Jones (pers. comm.) describes it as follows: ���The body is dark brown to brassy above, dark black lateral band, and brilliant white below, splashed with black blotches on the sides. Dorsal and caudal fins [are] dark olive-brown to reddish-brown, with the rays often deep-olive and with the nearly clear inter-radial membranes faintly flushed with brassy color; pectoral fins [are] yellowish with orange-red axils and very strong orange coloration at base; anal and pelvic fins [are] bright orange-red at the base with lessening coloration towards the rays. Cheeks and operculum with strong gilt reflections; strong orange coloration is found on the edges of the mouth (especially in males) with some blending into the upper mouth region. A lateral band is more strongly gilt than adjacent parts of body, thus often obscuring the lateral line. In females, the coloration is similar but less intense except for the orange coloration at the base of the paired fins that appears equally intense in both sexes. Males can be distinguished primarily by breeding tubercles on the top of the head.��� Distribution. The Red Hills Roach is confined to Six Bit Gulch and its tributary streams: Amber Creek, Horton Creek, Minnow Creek and Poor Man���s Gulch in Tuolumne Co. (Jones et al. 2002). Six Bit Gulch, during rainy periods, flows into the western arm of New Don Pedro Reservoir on the Tuolumne River. In July 2010, Roach were observed in three discontinuous wetted reaches of Horton Creek, which covered approximately 500 meters in total wetted length (P. Moyle, unpublished observations). However, only the lower reach, which extends about 200 meters upstream from the confluence with Six Bit Gulch, appeared to be perennial (as indicated by lush growth of sedges and other riparian vegetation). A natural fish barrier approximately 1.2 km upstream from the confluence likely inhibits Roach from accessing upper Horton Creek. Roach have also been observed in Six Bit Gulch where it is forded by Six Bit Ranch Road and in a pool in Roach Creek. Status. Red Hills Roach merit protection as a threatened or endangered subspecies and rate as an IUCN status of Critical Concern (Moyle et al. 2011, 2015). They have an extremely limited distribution in a very harsh environment and less than 1,000 individuals persist in isolated summer pools (Jones et al. 2002). Their persistence is threatened by fire, depleted stream flows, lack of protection on private lands, and most notably, from invasive fishes. While some habitat is protected in the Red Hills Area of Critical Environmental Concern, much is on private land along a road and remains unprotected. Red Hills Roach has persisted only because of its physiological capacity to endure the extreme environmental conditions found in these small exposed streams. Etymology. Both the common and subspecies name reflect the nature of the rocks through which these small creeks flow. The Red Hills region is named for the color of the soils created by weathering one of the largest outcroppings of serpentine rocks in the Sierra Nevada. Serpentine soils are high in magnesium and other minerals and create conditions in which few species can live. The landscape has a sparse but highly endemic flora (Anacker & Harrison 2012, Harrison 2013) and streams are small and largely unshaded., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 229-230, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Brown, L. R., Moyle, P. B., Bennett, W. A. & Quelvog, B. D. (1992) Implications of morphological variation among populations of California roach Lavinia symmetricus (Cyprinidae) for conservation policy. Biological Conservation, 62, 1 - 10.","Jones, W. J., Quelvog, B. D. & Bernardi, G. (2002) Morphological and genetic analysis of the Red Hills roach (Cyprinidae: Lavinia symmetricus). Conservation Genetics, 3, 261 - 276.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Moyle, P. B., Katz, J. V. E. & Quinones, R. M. (2011) Rapid decline of California's native inland fishes: a status assessment. Biological Conservation, 144, 2414 - 2423.","Moyle, P. B., Quinones, R. M., Katz, J. V. E. & Weaver, J. (2015) Fish Species of Special Concern in California. 3 rd edition. California Department of Fish and Wildlife, Sacramento Ca. 300 pp. https: // www. wildlife. ca. gov / Conservation / Fishes / Special-Concern","Anacker, B. L. & Harrison, S. P. (2012) Climate and the evolution of serpentine endemism in California. Evolutionary Ecology, 26, 1011 - 1023.","Harrison, S. (2013) Plant and Animal Endemism in California. University of California Press, Berkeley. 189 pp."]}

Lavinia and Hesperoleucus One conundrum within the Hitch���Roach species complex is uncertainty regarding whether species should be placed in one genus or two genera. While the complex was originally divided between Lavinia and Hesperoleucus, multiple examples of hybridization between the genera suggest they are more closely related than separate genera designations would normally indicate. For example, CA Roach are known to hybridize with Hitch in the Pajaro and Salinas rivers in Monterey Bay drainages (Miller 1945), in Coyote, Alameda, and Walnut Creeks in the San Francisco Bay region (Miller 1963; Leidy 1984, 2007, pers. comm.), and in Sacramento-San Joaquin drainages (Avise et al. 1975; Jones 2001). Hybrids were fertile in the Pajaro River; Avise et al. (1975) found 8% of Hitch examined to be F 1 hybrids and 5% to be backcrossed. Since it is unusual for two genera to be interfertile and because other endemic species in California do not appear to exhibit this behavior (Avise et al. 1975; Avise & Ayala 1976; Moyle & Massingill 1981; Moyle 2002), it has been proposed these two species belonged in a single genus. Aguilar & Jones (2009) found the two genera are closely related mtDNA lineages that could be considered to be one genus. Sch��nhuth et al. (2012), however, noted that while their mtDNA analyses supported this close relationship, their limited nuclear DNA results did not. Our recent study (Baumsteiger et al. 2017) found the strongest genomic signal coincided with splitting samples into two genera. We discovered, however, that samples from the Pit River (Northern Roach) were a confounding factor, having mixed historical ancestry with both genera. It was estimated in one analysis that approximately 80% of the Northern Roach genome clustered with Hitch whereas only 20% clustered with CA Roach. But all subsequent analyses found Northern Roach to be a distinct species, on par with currently identified species. Thus Northern Roach appears to be the result of an ancient hybrid speciation event (Mallet 2007) between ancestral Hitch and CA Roach but it is currently genomically different enough to warrant species status. It is our conclusion that despite hybridization between individuals in the different genera, the distinctiveness of each genus is still strongly maintained and prevalent throughout the genomes interrogated. If results were more admixed, the argument for a single genus would be more plausible. We recognize admixture between genera is uncommon but note that not all mutations lead to post-zygotic barriers (Abbott et al. 2013). Additionally, hybridization has been proposed between Roach/Hitch and other members of the subfamily Leuciscinae including Speckled Dace (Rhinichthys osculus), Arroyo Chub (Gila orcuttii), and the now extinct Thicktail Chub (Gila crassicauda) (Greenfield & Deckert 1973; Hopkirk 1973). Thus the frequency of hybridization suggests reproductive barriers are poorly developed in the endemic minnows of California. Secondly, although Northern Roach represent an ancient hybridization event and cluster primarily with individuals from the Lavinia genus (~80% but only in the admixture analysis), we cannot definitively say they are Lavinia. Northern Roach still strongly resemble CA Roach in appearance (see below), but this may simply be the result of natural selection for the CA Roach morphotype. Future phylogenetic analyses between all genera within the family may provide the solution but it remains, as yet, unavailable. Thus, without universal agreement between genomic analyses or improved criteria for taxonomically delineating hybrids, we default to the original genus assignment by Snyder (1913). Therefore, the species of Hesperoleucus are as described below., Published as part of Baumsteiger, Jason & Moyle, Peter B., 2019, A reappraisal of the California Roach / Hitch (Cypriniformes, Cyprinidae, Hesperoleucus / Lavinia) species complex, pp. 221-240 in Zootaxa 4543 (2) on pages 223-224, DOI: 10.11646/zootaxa.4543.2.3, http://zenodo.org/record/2617771, {"references":["Miller, R. R. (1945) The status of Lavinia ardesiaca, a cyprinid fish from the Pajaro-Salinas River basin, California. Copeia, 1945, 197 - 204.","Miller, R. R. (1963) Synonymy, characters, and variation of Gila crassicauda, a rare Californian minnow, with an account of its hybridization with Lavinia exilicauda. California Fish and Game, 49, 20 - 29.","Leidy, R. A. (1984) Distribution and ecology of stream fishes in the San Francisco Bay drainage, Hilgardia, 52 (8), 1 - 175.","Leidy, R. A. (2007) Ecology, Assemblage Structure, Distribution, and Status of Fishes in Streams Tributary to the San Francisco Estuary, California. San Francisco Estuary Institute, San Francisco. 194 pp.","Avise, J. C., Smith, J. J. & Ayala, F. J. (1975) Adaptive differentiation with little genic change between two native California minnows. Evolution, 29, 411 - 426.","Jones, W. J. (2001) DNA Sequence Divergence and Speciation in Two California Minnows (Cyprinidae: Lavinia exilicauda and L. symmetricus). PhD Dissertation, University of California, Santa Cruz. Ca. 150 pp.","Avise, J. C. & Ayala, F. J. (1976) Genetic differentiation in speciose versus depauperate phylads: evidence from the California minnows. Evolution, 30, 46 - 58.","Moyle, P. B. & Massingill, M. (1981) Hybridization between hitch, Lavinia exilicauda, and Sacramento blackfish, Orthodon microlepidotus, in San Luis Reservoir, California. California Fish and Game, 67, 196 - 198.","Moyle, P. B. (2002) Inland Fishes of California. University of California Press, Berkeley, California. 502 pp.","Aguilar, A. & Jones, W. J. (2009) Nuclear and mitochondrial diversification in two native California minnows: insights into taxonomic identity and regional phylogeography. Molecular Phylogenetics and Evolution, 51, 373 - 381.","Schonhuth, S., Hillis, D. M., Neely, D. A., Lozano-Vilano, L., Perdices, A. & Mayden, R. L. (2012) Phylogeny, diversity, and species delimitation of the North American round-nosed minnows (Teleostei: Dionda), as inferred from mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution, 62, 427 - 446.","Baumsteiger, J., Moyle, P. B., Aguilar, A., O'Rourke, S. M. & Miller, M. R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PloS One, 12 (12), e 0189417.","Mallet, J. (2007) Hybrid speciation. Nature, 446 (7133), 279.","Abbott, R., Albach, D., Ansell, S., Arntzen, J. W., Baird, S. J., Bierne, N., Boughman, J., Brelsford, A., Buerkle, C. A., Buggs, R. & Butlin, R. K. (2013) Hybridization and speciation. Journal of Evolutionary Biology, 26, 229 - 246.","Greenfield, D. W. & Deckert, G. D. (1973). Introgressive hybridization between Gila orcutti and Hesperoleucus symmetricus (Pisces: Cyprinidae) in the Cuyama River Basin, California: II. Ecological aspects. Copeia, 1973, 417 - 427.","Hopkirk, J. D. (1973) Endemism in fishes of the Clear Lake region of central California. University of California Publications in Zoology, 96, 1 - 160.","Snyder, J. O. (1913) The fishes of the streams tributary to Monterey Bay, California. Bulletin, U. S. Bureau of Fisheries, 32, 49 - 72."]}