6,835 research outputs found
Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA
No full textMoyle, Peter B., Buckmaster, Nicholas, Su, Yingxin (2023): Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA. Zootaxa 5249 (5): 501-539, DOI: 10.11646/zootaxa.5249.5.1, URL: http://dx.doi.org/10.11646/zootaxa.5249.5.
FIGURE 4 in Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA
No full textFIGURE 4. Holotypes of new taxa of Rhinichthys described in this paper, A. Santa Ana Speckled Dace (R. gabrieleno WFB 3498. B. Long Valley Speckled Dace (R. nevadensis caldera) WFB 5000, C. Warner Speckled Dace (R. klamathensis goyatoka) WFB 122-10-44, D. Sacramento Speckled Dace (R. klamathensis achomawi) WFB 3171.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 page 515, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/770135
Rhinichthys nevadensis subsp. caldera Moyle & Buckmaster & Su 2023, new subspecies
No full textRhinichthys 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/770135
Interview with Nicholas Christopher, author of Somewhere in the Night: Film Noir and the American City
No full textInterview with Nicholas Christopher, author of Somewhere in the Night: Film Noir and the American Cit
Rhinichthys nevadensis subsp. robustus Moyle & Buckmaster & Su 2023, new combination
No full textRhinichthys 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 <8 cm SL) active fish with blotches on the side that can merge to become a stripe leading to a dark band around the eye. The caudal peduncle is thick, about half the body depth, while the body is subcylindrical (robust). The head is bluntly pointed, with a tiny sub-terminal mouth. Fins are small (the dorsal fin usually has 8 rays, the anal 7 rays). Presence of tiny maxillary barbels at the corners of the mouth is variable as is the presence of a frenum. It is distinguished from the other two subspecies of 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/770135
Rhinichthys klamathensis subsp. goyatoka Moyle & Buckmaster & Su 2023, new subspecies
No full textRhinichthys 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 (<10 cm SL); classic Speckled Dace body shape: thick caudal peduncle, robust (subcylindrical) body, small fins, and bluntly pointed snout with subterminal mouth. Maxillary barbels and frenum usually present. 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/770135
Rhinichthys klamathensis subsp. klamathensis Moyle, Buckmaster & Su, 2023, new combination
No full textRhinichthys 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 (<1 million years), including Jenny Creek as a distinct population. Overall, analysis of mtDNA data of dace from their entire range showed that dace from the Klamath, Sacramento, and Warner basins share much of the same lineage; together they form a separate lineage within the Northwest clade of Smith 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/770135
Rhinichthys klamathensis subsp. achomawi Moyle & Buckmaster & Su 2023, new subspecies
No full textRhinichthys 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 <8 cm SL, occasionally to 11 cm SL), highly variable species distinguished by a thick caudal peduncle, a small subterminal mouth, a pointed snout, and small scales (47–89 in lateral line). The origin of the dorsal fin (6–9 rays, usually 8) is well behind that of the pelvic fins. The anal fin normally has 7 rays (6–8). The pharyngeal teeth (1,4-4,1 or 2,4-4,2) are strongly hooked and have only a slight grinding surface Color is highly variable but most fish over 3 cm SL have dark speckles on the sides and back, dark blotches on the sides that often coalesce to resemble a dark lateral band, and a stripe on the head that runs through the snout. The background color on the back and sides is dusky yellow to dark olive, with the belly yellowish to whitish. The bases of the fins of both sexes turn orange to red during breeding and males often have red snouts and lips as well. Presence of maxillary barbels and a frenum is variable. Rutter (1908:140) noted that Sacramento dace were quite variable in their characteristics and found that Sacramento dace from 12 localities (n=94) had 49–77 scales in the lateral line, 7–9 dorsal rays, and 6–8 anal rays. Cornelius (1969) using data from six localities, counted 54–82 scales on the lateral line (mean 70, n = 123); other counts: scale rows above the lateral line (12–20, mean 16, N = 124) scale rows below lateral line (10–19, mean 14, n = 121), vertebrae (34–39, mean 37, n=99), dorsal-fin rays (7–9, mean 8, n=131), anal-fin rays (5–8, mean 7, n =129). See also Table 3. 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/770135
Resurrecting the Author
No full textPresentation of Nicholas Wolterstorff\u27s Paper Resurrecting the Author with time after for questions beginning at 18:00
FIGURE 2 in Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA
No full textFIGURE 2. Phylogenetic relationships among Speckled Dace populations in this study, modified from Su et al (2022). The molecular phylogeny was generated by an ML coalescent model in SVDQuartets with 100 bootstraps, using RAD sequencing data. Numbers on each lineage refer to the percentage of bootstrap runs (out of 100) that produced the same result. Details can be found in Su et al. (2022). The lineage names refer to major drainages from which our Speckled Dace samples were collected from California, as well as from major basins outside of California. We did not examine the latter findings in detail; they probably represent multiple undescribed species/subspecies. The capital letters refer to lineages identified as species in this paper. A. Desert Speckled Dace (R. nevadensis), B. Western Speckled Dace (R. klamathensis), C. Santa Ana Speckled Dace (R. gabrielino). Tui Chub (Siphatales bicolor) and Relict Dace (Relictus solitarius) were outgroups used to root the phylogeny because they are both western cyprinids distantly related to Rhinichthys (Schönhuth et al. 2012).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 page 512, DOI: 10.11646/zootaxa.5249.5.1, http://zenodo.org/record/770135
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