NATIVE PISCIFAUNA OF UTAH LAKE

A. GAYLON COOK

BIOCONSERVATION INTERNATIONAL 3306 N CANYON RD PROVO UT 84604-4548

Running Head: NATIVE PISCIFAUNA OF UTAH LAKE NATIVE PISCIFAUNA OF UTAH LAKE

A. Gaylon Cook'

ABSTRACT.— A thorough search of the descriptive and systematic ichthyological literature disclosed the piscine specimens from Utah Lake, and manifested an elaborate web of supposed synonyms for which there were nomenclatural as well as species-assignment errors. In this review, the characteristics of piscine examples from Utah Lake, and forms that have been considered to be their conspecifics, are delineated and contrasted; deductions about valid synonyms are made. A careful comparison of the morphologic descriptions of exemplars of Utah lake forms and their putative conspecifics requires that the synonymies of several taxa be pared. By providing a comprehensive bibliography, and furnishing winnowed synonymies of Utah Lake forms that signal which forms to reexamine, this paper establishes a basis for needed chemical taxonomic analyses such as DNA-DNA hybridization tests. Also, the detailed descriptions of taxa available herein constitute a resource that can be used for such purposes as distinguishing species in periodic ichthyological surveys of Utah Lake or identifying taxa (e.g., suckers) during endangered species recovery-plan activities.

Key Words: piscifauna, ichthyofauna, Utah Lake, fish, fishes, taxonomy, systematics, synonymy, native.

Utah Lake, closely bordered by the city of Provo, is a 38,800-ha body of fresh water in central Utah measuring 23.5 by 38.0 km (Carter 1969: 2, Radant and Sakaguchi 1981: derived from Fig. 1). It has an average depth of 2.9 m and a maximal depth of 4.2 m (Radant and Sakaguchi 1981: 2). Formerly it collected a flow of 740 million m3/yr (Arnold 1960: 11-17). Jordan and Gilbert (1881) provided the first report on the ichthyofauna of Utah Lake; 1 form collected near Evanston, Wyoming was included. Although Jordan

(1891) furnished an account of the fishes of Salt Lake Valley and Utah Valley, whose combined areas he considered to constitute "Salt Lake Basin", the next treatise specifically treating the piscifauna of Utah Lake was Hatton's (1932) review. The most recent document on this subject was an article by Heckmann et al.

(1981) whose cursory treatment of native fishes consisted principally of an unreferenced annotated list. La

'Bioconservation International, Provo, Utah 84604-4548. GREAT BASIN NATURALIST 2

Rivers' (1962) reference contains synonymic lists for most of the ichthyofauna of Utah Lake. No work has comprehensively characterized these fishes or unsnarled and expounded their synonymies. Many publications

(see below) have treated 1 piscine taxon from Utah Lake. In totality, they evince multiple nomenclatural errors, and remarkably extensive synonymies for most taxa due partially to age-related differential morphologies. Herein, the chronologies and explanations of reported (and often inordinate) synonymies, are based on a perusal of a plethora of descriptive or taxonomic documents pertaining to native Utah Lake fishes or putative synonyms. All were scrutinized and compared to reveal corroborations, or discrepancies and contradictions. Piscine taxonomy has been affected most by the massive lumping of species by Jordan and

Evermann (1896a,b, 1898a,b). That action's impact persists. Of the 11 piscine forms from Utah Lake that were considered to be species by Robins et al. (1991), 8 were regarded as polytypic by Mayden et al. (1992: table 1). Ignorance about the phylogenetic relationships of Utah Lake fishes is underscored by the dispute about the number of sucker species present (see below). Significantly, it is not known whether certain piscine taxa of Utah Lake constitute endemic species or subspecies that would be worthy of endangered status.

The principal aim of this review is to establish a sound basis for defining and delimiting as precisely as possible the native piscifauna of Utah Lake. It is hoped that it will lay the groundwork for further systematic investigations. Taxonomic studies employing DNA-DNA hybridization analyses, as well as those that might utilize biochemistry, immunology, serology, protein sequencing, or comparative anatomy, need to be performed on Utah Lake's native fishes and their putative conspecifics. Besides being quick, DNA hybridization offers the most precise quantitative comparisons (Sibley and Alquist 1990: 3). It is also anticipated that the characterizations that are presented here will assist ichthyologists in identifying Utah

Lake fishes. For example, descriptive data on catostomids can be used by field biologists with the Utah

Department of Natural Resources, Division of Wildlife Resources in their effort to identify endangered adult

June Suckers (Chasmistes Bow), and distinguish them from Webug Suckers (Chasmistes fecundus) and Utah

Suckers (Catostomus ardens) during spawning-season sampling and reproductive facilitation in lower Provo

River. Suckers are frequently misidentified by these biologists during reproductive surveys that are NATIVE PISCIFAUNA OF UTAH LAKE 3 performed as a part of the June Sucker recovery plan (Charles W. Thompson, personal communication). The temporally-related comparative incidence of the fishes of Utah Lake will be treated in a future paper.

Textual and tabular notes of the Utah Lake Fish Management Advisory Team (1994: 16, table 1) indicating that Cope and Yarrow (1876) identified examples of 4 piscine species from Utah Lake, and that

1 of them was Gila nigrescens, are in error.

ANNOTATED SYNONYMIES AND DESCRIPTIONS OF TAXA

CLASS OSTEICHTHYES

ORDER CYPRINIFORMES

Family Cyprinidae

Leatherside Chub Gila copei (Cope's Gila: Gila is a river near the border of USA and Mexico; copei refers to Professor Edward Drinker Cope, who as a leader of the Wheeler Survey, collected and identified many animals in the western USA in the 1870s). The description of Tigoma gracilis, taken at 41° N latitude

(Girard 1857a: 26, c: 166, 206-207, 1858b: 293), has been interpreted (Jordan 1891, Jordan and Evermann

1896a: 236, Jordan et al. 1930: 119, Hayes 1935: 67) to indicate that this form was a synonym. Girard

(1858b) did not equate his T. gracilis with Gila gracilis of Baird and Girard (1854a) but Cope and Yarrow

(1876: 665) and Jordan (1878a: 424) later did. Jordan (1887b: 818-819) considered G. gracilis and Phoxinus

(=Tigoma) gracilis to be separate. Jordan and Evermann (1896a: 227-228) and Jordan et al. (1930: 114) synonymized G. gracilis, which had a short, deep body and dorsal and anal fins bearing 3 rudimentary spiny rays, with Gila robusta. In contrast to G. gracilis, the Leatherside Chub from Utah's Provo and Spanish

Fork Rivers had next to the slenderest body (body depth / body length [BD/BL] = 22.4 ± 1.5% [± 1 SD from the mean]; n = 55) and next to the slimmest caudal peduncle (depth = 11.0 ± 0.5% of BL En = 54]; length = 23.3 ± 1.2% of BL = 551) (see Girard 1857a,c, 1858b, Jordan and Gilbert 1882: 238-239,

Jouy 1882, and Jordan and Evermann 1896a: 236) of the 6 cyprinids of Utah Lake and its affluents (Hayes

1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37). Jordan (1891) reported that the 2 types of the vaguely described T. gracilis had been lost; Jordan et al. (1930: 119) noted that this form was unidentifiable. GREAT BASIN NATURALIST 4

There is no evidence to support Jordan's (1887a) report that Platygobio gracilis exists in Salt Lake

Basin. Jordan and Gilbert (1881) described examples of Squalius copei from tributaries of Bear River near

Evanston, Wyoming, affirming that Hybopsis egregius of Cope (1872a) and Gila egregia of Cope and

Yarrow (1876: 662) were synonyms of it but that Tigoma egregia of Girard (1857a: 25, 1858b: 291-292) was not because the number of scales along its lateral line (Su = 66) (see also Jordan 1891; Jordan and

Gilbert [1882: 236] indicated that Su = 65) was too low. However, Cope (1872a) had regarded H. egregius and T. egregia as synonymous. Accordingly, Jordan and Gilbert (1882: 238-239) did not include the sketchily described H. egregius as a synonym of S. copei either. Jordan (1887b: 819) regarded Phoxinus (=

Tigoma) egregius and Phoxinus (= Squalius) copei as distinct forms. Jordan (1891) excluded Cope and

Yarrow's (1876) G. egregia from "Loma, Rio Grande, Colo" as a synonym of Leuciscus copei, while including G. egregia from Utah's Beaver River. Other synonyms of the Leatherside Chub are Alburnellus? sp. from Utah Lake (Cope 1876) and Squalius aliciae (Phoxinus aliciae of Jordan [1887b]) from Utah Lake

(Jordan and Gilbert 1882: 238, Jouy 1882) or Provo River near that lake (Jordan and Evermann 1896a: 236, b: 248). Although Jordan and Evermann (1896a) revealed with a list of 3 synonyms that copei had priority

(the types of T. gracilis had been lost and gracilis was preoccupied in Leuciscus), they named the species

L. aliciae after the wife of a biologist with whom Jordan coauthored a work (Jordan and Jouy 1882) rather than after Cope, but then reversed themselves (Jordan et al. 1930: 119), calling the taxon Cheonda copei.

Later, it was referenced as Richardsonius copei (Hayes 1935: 67, Tanner 1936), Snyderichthys copei (Bailey et al. 1960: 17), and finally Gila copei (Sigler and Miller 1963: 74-76, Robins et al. 1991: 20).

This species had a rounded snout, equal jaws, and a rather small, low, terminal, and oblique mouth with irregular, uncinate teeth (Jordan and Gilbert 1882: 238-239, Jouy 1882, Jordan and Evermann 1896a:

236, Hayes 1935: 68). Maxillae extended caudad to a point ventrad to the rostral border of the eyes (Girard

1857a: 26, 1858b: 293, Jordan and Gilbert 1882, Jouy 1882, Hayes 1935). Jordan and Evermann's (1896a) statement that maxillae continued slightly beyond that point was not grounded in evidence. The broad head

(width f.-- 56-60% of length) had a slightly convex interorbital span which in S. aliciae (Jordan and Gilbert NATIVE PISCIFAUNA OF UTAH LAKE 5

1882, Jouy 1882) = 8.2% of BL, and in R. copei (Hayes 1935: derived from tables 22-23 and 27-28) = 8.7

± 0.5% of BL, a width > eye diameter which = 5.8 ± 0.8% of BL. Eye diameter / snout length (ED/SL) averaged 0.85 (SL/BL = 6.9 ± 0.4%) in R. copei and = 1 in S. copei; ED / head length (HL) = 20% in

T. gracilis, 25% in S. aliciae whose total length (TL) = 67 mm, 25.7% in R. copei whose TL = 85 mm

(BL = 69.0 ± 13.6 mm), and 25-29% in S. copei whose TL = 152 mm (Girard 1857a, 1858b, Jordan and

Gilbert 1882, Jouy 1882, Hayes 1935). Girard (1857a) declared that the HL/TL of T. gracilis...--.- 22%, but also (Girard 1857c: 206-207, 1858b: 292-293) noted that it was > that (slightly < 14) of T. lineata. HL/BL

= 22.8 ± 1.2% (n = 55) in R. copei, 23.1 or 23.9% in S. aliciae, and 23.5% in S. copei (Jordan and

Gilbert 1882, Jouy 1882, Hayes 1935). The HL/TL values of =22% and slightly < '4 in gracilis were appreciably > the HL/BL values of 22.8-23.9% in supposed conspecifics. In R. copei, for example, mean

HL/TL = 18.5%. Since both relative eye diameter (ED/BL) and head length (HL/BL) in Hayes' (1935)

R. copei were negatively correlated with BL (r = -0.852 and -0.634, respectively; df = 53, P < 0.001),

ED and HL were positively correlated (r = 0.765, P < 0.001). Thus, the great difference between the

ED/HL of gracilis (20%) and other forms (25-29%), may be taxonomically meaningful. The atypical HL and ED of gracilis suggest that it may not have been a Leatherside Chub; it will not be considered further.

The lateral line was complete and slightly deflexed (Jordan and Gilbert 1882: 238-239, Jouy 1882).

Su equaled 77 in Alburnellus? sp. from Utah Lake and Utah's Beaver River (Cope 1876) and G. egregia from Beaver River (Cope and Yarrow 1876: 662), 80 in S. aliciae from Utah Lake (Jouy 1882) and S. copei from Evanston, Wyoming (Jordan and Gilbert 1882: 238-239), and 81.3 ± 4.2 in Hayes' (1935) R. copei.

There were 18 longitudinal rows of scales above the lateral line (Sa = 18) in Alburnellus? sp., G. egregia, and S. aliciae, 18.1 ± 1.3 in R. copei, and 19 in S. copei. Below the lateral line, there were 9 scale rows

(Si, = 9) in Alburnellus? sp. and G. egregia, 11.7 ± 0.8 in R. copei, 12 in S. copei, and 15 in S. aliciae.

The mode in the number of dorsal-fin and anal-fin rays was 8; the dorsal fin originates slightly caudad to the origin of pelvic fins (P2) (Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882, Jouy 1882,

Jordan and Evermann 1896a: 236, Hayes 1935). Predorsal length was 53.2 ± 1.5% of BL, and the distance GREAT BASIN NATURALIST 6

from the tip of the snout to the origin of P2 was 47.7 ± 1.6% (Hayes 1935). Pectoral fins (P,), although slender and rather long (21.0 ± 2.0% of BL in R. copei and 23.9% in S. aliciae), fell far short of reaching

the squat P2 (length = 15.3 ± 1.2% of BL) which did not reach the vent caudally (Jordan and Gilbert 1882,

Jouy 1882, Jordan and Evermann 1896a, Hayes 1935). In R. copei, the basal lengths of the dorsal and anal

fins = 11.9 ± 0.8% of BL and 11.0 ± 0.7% of BL, respectively (Hayes 1935). The height of the dorsal

fin = 20.9% in S. aliciae and 21.5 ± 1.5% in R. copei; anal-fin depth = 16.4% in S. aliciae and 18.5 ±

1.4% in R. copei. Jordan and Gilbert (1882) remarked that S. copei had a bluish olive dorsum with dark

stippling, and dusky sides. Jouy (1882) recounted that S. aliciae had a smoky dorsum, pinldsh venter, steely

blue lateral band, and silvery cheeks. Hayes (1935: 68) stated that the dorsum was olivaceous anthracine

(bluish black) and the sides were dusky. Fin axils were scarlet (Hayes 1935), crimson (Jordan 1891), or red

(Jordan and Gilbert 1882, Jordan and Evermann 1896), at least in males. Hayes (1935) affirmed that

specimens from alkaline or turbid waters were more pallid than those from clear streams.

Utah Chub Siboma atraria (blackish siboma: siboma is apparently a euphonic North American Indian word

[see Jordan and Evermann 1896a: 229]; Latin atra, black; -anus, a suffix meaning pertaining to). Putative

binomial synonyms include Tigoma obesa, T. lineata, and Siboma atraria (Girard 1857a: 25-26, c: 206, 208,

1858b: 290-293, 297-298); Tigoma squamata (Gill 1862b, 1876); Protoporus domninus, Hybopsis bivittatus,

S. atraria, and Myloleucus pulverulentus (see Hayes [1935]) (Cope 1872b); S. atraria and Hybopsis

timpanogensis (Cope 1876); H. timpanogensis, S. atraria, M. pulverulentus, and Orthodon microlepidotus

(Cope and Yarrow 1876: 654-655, 667-669, 681); O. microlepidotus, Alburnops bivittatus,

A. timpanogensis, P. domninus, Gila obesa, G. lineata, S. atraria, and M. pulverulentus (Jordan 1878a),

Squalius cruoreus, S. rhomaleus, and S. atrarius (Jordan and Gilbert 1881); 0. microlepidotus, Minnilus

bivittatus, M. timpanogensis, P. domninus, Squalius cruoreus, S. lineatus, S. obesus, S. rhomaleus,

S. squamatus, S. atrarius, and Leucus obesus (includes M. pulverulentus) (Jordan and Gilbert 1882: 152,

195-196, 204, 234-237, 240-241, 245, 886-887); S. atrarius (Jordan 1886b); S. atrarius and S. rhomaleus

(Jordan 1887a), 0. microlepidotus, Notropis ? bivittatus, N. ? domninus, N. ? timpanogensis, Phoxinus NATIVE PISCIFAUNA OF UTAH LAKE 7 ctuoreus, P. lineatus, P. obesus, P. atrarius, and P. squamatus (Jordan 1887b: 808, 814-815, 819);

Leuciscus atrarius (Jordan 1891, Evermann 1893); L. lineatus and Rutilus symmetricus (comprises

M. pulverulentus) (Jordan and Evermann 1896a: 232-233, 245-246, b: 248, 250), L. lineatus (Jordan and

Evermann 1902: 70-71), and eventually Gila atraria (La Rivers and Trelease 1952: 116) (see below).

The record of Cope and Yarrow's (1876: 681) Orthodon microlepidotus indicates that the form which

Mormon settlers called "chub" was abundant in Provo River, and Utah Lake where it was collected in July

1872. 0. microlepidotus, which reportedly spawned in muddy streams during May and June, was "terribly destructive to" piscine eggs and fry, and was a "fair table fish." Substantiating that this form was the Utah

Chub, Jordan and Gilbert (1881) declared that the "'chub' is very abundant in Utah Lake(,) ...exceedingly destructive to the young" trout, and was christened "devilish chub" by Peter Madsen, a Danish Mormon immigrant and expert fisherman (see Carter 1969). Residents of Utah Valley have consistently referred to this species as "chub" (Carter 1969). Jordan and Gilbert (1882: 152) noted that 0. microlepidotus inhabited the Great Basin of Utah as well as California rivers, but they morphologically described Gila microlepidota of California (Ayres 1854) whose Su = 105-110, S. = 21-24, and Si, = 12. Cope (Cope and Yarrow 1876) declined to vouch for the locality of the lot of specimens supposedly taken from Utah Lake because he considered 0. microlepidotus to exclusively reside on the Pacific slope. Cope and Yarrow (1876) neglected to mention that the description of the "chub" might have become associated with the wrong specimen label.

Girard (1857a: 25-26) assigned the generic name Tigoma to obesa from Salt Lake Valley, and then to 4 other forms, including lineata. This publication was a valid one according to Article 8 of the

International Code of Zoological Nomenclature (Ride et al. 1985), most importantly because numerous identical copies of it were simultaneously reproduced. Article 12 of that code stipulated that a name published before 1931 must have been accompanied by a description, definition, or 1 of 8 "indications", the 5th of which (see Article 16) manifests that a new generic name be cited in combination with 1 available specific epithets. Tigoma and several of its members were described by Girard (1857c: 205), as was Siboma atraria, an appellation for the Utah Chub. Cope and Yarrow (1876: 667-668), Jordan and GREAT BASIN NATURALIST 8

Gilbert (1882: 241, 886-887), Jordan and Evermann (1896a: 232-233, b: 248), and Robins et al. (1991: 20) did not cite Girard (1857a) but alluded to Girard (1857c) with the date, 1856, the year of the proceedings of the Academy of Natural Sciences of Philadelphia that are in volume 8, rather than the year of that volume's publication appearing on its title page (1857). Robins et al. (1991: 109) contended that pp. 161-258 of volume 8 were issued prior to 28 December 1856. Jordan (1891) named the Utah Chub Leuciscus atrarius, to which T. obesa was synonymized. Jordan and Evermann (1896a) signified that because obesus

(masculine of obesa, the 6th taxon treated by Girard [1857c] on p. 206) was preoccupied in the genus

Leuciscus, Jordan (1891) had made S. atraria, described by Girard (1857c) on p. 208, the nominal form.

In substance, Jordan (1891), the "first reviser" (Ride et al. 1985: article 24) of the nomenclature of the Utah

Chub gave obesa precedence over other synonyms. Jordan (1891) and Evermann (1893) reexamined specimens of S. cruoreus and concluded that they were juveniles conspecific with adult examples of L. atrarius, and also synonymous with Squalius (= Tigoma) squamatus and S. rhomaleus. Jordan (1891) supposed that Hybopsis (=Minnilus) bivittatus and H. timpanogensis were also conspecific. Evermann (1893) surmised that Gila egregia (Cope and Yarrow 1876: 662) from Utah's Beaver River was additionally a synonym, but this form = G. copei (see above). Jordan and Evermann (1896a) decided that T. lineata, whose provenance was unknown and type had been lost, was a Utah Chub and made it the nominal form; lineata was the 8th species described by Girard (1857c) on p. 206. Snyder (1922) stated that only 1 pharyngeal-arch specimen of lineata remained, bearing the U.S. National Museum Catalog No. 2783. Girard

(1858b: 292-293) had noted that 2783 was the label number for the teeth of lineata, but that the catalog number for a type series of 6 specimens was 229. Snyder (1922), who noticed that pharyngeal teeth of lineata resembled those of Richardsonius hydrophlox, and that its body shape was reminiscent of R. egregius , if not R. hydrophlox, called the Utah Chub Richardsonius atrarius. However, A (the number of rays in the anal fin) = 11 in R. hydrophlox (Cope 1872b) but only 8 in lineata (Girard 1858b). The traits of lineata fit those of Gila copei which comprises G. egregia from Utah's Beaver River (see above). Instead of having a "short and compact", "robust", "very robust", "stout", "rather stout", "somewhat elongated", or NATIVE PISCIFAUNA OF UTAH LAKE 9

"moderately elongate" body like its putative synonyms, lineata uniquely had an elongate body that tapered caudally (Girard 1857a: 25-26, c: 206, 208, 1858b: 290-293, 297-298, Jordan and Gilbert 1882: 195-196,

204, 234-237, 240-241). Any persisting exemplars of lineata (have all 6 been lost?) ought to be reexamined.

Jordan etal. (1930: 119) reiterated Snyder's (1922) finding that lineata was not identifiable, and again made atraria the nominal form of all synonyms of the Utah Chub except Myloleucus pulverulentus. They placed the latter (referenced with the feminine pulverulenta) and atraria in the genus Tigoma. Based on the precedence of obesa and the fact that it was not preoccupied in the genus Tigoma, the establishment of atraria as the nominal form was a mistake. obesa was not even listed as its synonym (or any synonym).

Tigoma obesa (Girard 1857a: 25, c: 206, 1858b: 290-291), considered by Jordan (1891), Jordan and

Evermann (1896: 232-233), and Hayes (1935: 40) to be a synonym of the Utah Chub, was likely

R. hydrophlox. In seeming distinction, obesa was "short and compact" whereas R. hydrophlox, whose

BD/BL reached 14 (see below), was neither slender nor especially squat. Significantly, however, obesa's anal fin was larger than its dorsal fin and possessed more rays (A = 9) than the anal fin of putative synonyms (D [the number of rays in the dorsal fin] = 8 in obesa) (Girard 1858b, Jordan and Gilbert 1882:

237). In hydrophlox (see below), A = a mode of 10 or 11 but equaled 9 in multiple 1934 examples of it

(Hayes 1935: tables 17-20). In the Utah Chub, the anal fin is smaller than the dorsal. The basal length of the anal fin = 7.1-7.7% of TL in atraria (Girard 1858b: 297-298), and the basal length and depth of the anal fin = 8.3% and 14.4% of TL (10.4 + 0.5% and 18.0 ± 1.3% of BL), respectively, in 16 specimens collected from Utah Lake in 1934, and 9.1% and 13.0% of TL (11.4% and 16.2% of BL), respectively, in squamata; in contrast, the basal length and height of the dorsal fin = 10.7% and 18.1% of TL (13.3 ±

0.8% and 22.6 ± 1.0% of BL), respectively, in 1934 specimens, and 11.1% and 15.0% of TL (13.9% and 18.8% of BL), respectively, in squamata (Gill 1876, Hayes 1935: derived from table 2). In examples of the Utah Chub from Utah Lake (timpanogensis, rhomaleus, cruoreus, and atraria (in part), A = 8 and

D had a mode of 9; in putative synonyms collected elsewhere (bivittatus, squamata, atraria [in part], and domninus), A = 7-8 (mode of 8) and D = 8-9, except for atraria in which A = 8-9 and D = 8-10 (Girard GREAT BASIN NATURALIST 10

1858b: 292-293, 297-298, Cope 1872b, 1876, Cope and Yarrow 1876: 654-655, 667-668, Gill 1876, Jordan and Gilbert 1881, 1882: 195-196, 204, 234-237, 240-241, 886-887, Snyder 1922, Hayes 1935). According to Sigler and Sigler (1987: 163-166), the range of the Utah Chub is limited to the watersheds of ancient Lake

Bonneville and Snake River. Cope and Yarrow's (1876: 667-668) curaria, which comprised non-Utah specimens from Denver, Colorado as well as Zuni and Little Colorado (Colorado Chiquito) Rivers of western New Mexico, was thus, likely polytypic, accounting for the high fin-ray counts of some exemplars.

Caudally, P2 of obesa did not come close to the vent (Girard 1858b: 290-291). In Utah Chubs, P2 approach or reach the vent and P1 extend 50-60% of the distance to P2 (Girard 1858b: 297-298, Cope 1872b, 1876,

Cope and Yarrow 1876: 654-655, Jordan and Gilbert 1881, 1882: 195-1%, 234-235, 240, Jordan and

Evermann 1896a: 232-233). Dorsally juxtaposed to the lateral line of obesa was a reddish lateral streak

(Girard 1857a, 1858b: 290-291). Although its body was squat, obesa had a slender caudal peduncle. The

Utah Chub's caudal peduncle was deeper than average (11.6 ± 0.7% of BL En = 16]) for the cyprinids of

Utah Lake and its tributaries (Jordan and Gilbert 1881, 1882: 237, 240-241, 886-887, Hayes 1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37); however, Carbine (1936: derived from table 5) contrastively reported that caudal-peduncle depth in 16 Utah Chubs that were collected during 1934-1935, including 5 15 from Utah Lake and 1 from Provo River, = 9.5 ± 0.5% of BL. The caudal extent of P2, comparative lengths of the anal and dorsal fins, willowy caudal peduncle, reddish vitta, and D are all traits of obesa which fit R. hydrophlox (see below). Dr. Charles H. Gilbert (Jordan 1891) compared obesa and cruoreus, and judged that they were equal, but his examination of the 3 specimens of obesa collected in 1852, bloated due to the poor quality of preservative alcohol, may have been hampered (Girard 1858b).

La Rivers and Trelease (1952: 116) subsumed atraria within the genus Gila. Gila, which Baird and

Girard (1854a,b) assigned to 5 species, had priority over Tigoma or Siboma, both initially presented by

Girard (1857a: 25-26) and described by Girard (1857c: 205, 208). Coburn and Cavender (1992: 344) did not discover any synapomorphies in osteological or scale traits within members of the currently recognized

(Robins et al. 1991) Gila genus to validate it. Derived characters that Gila spp. share with several other NATIVE PISCIFAUNA OF UTAH LAKE 11 putative genera confirm Barbour and Miller's (1978: 61-62) inference that Gila is paraphyletic. Coburn and

Cavender's (1992: fig. 3) analyses indicate that supposed members of Gila belong to 5 separate taxonomic categories. Placing the Utah Chub in the genus Gila is misleading. This practice insinuates nonexistent phylogenetic relationships with 15 supposed congeners (see Robins et al. 1991: 20). With the exception of

G. crassicauda, other supposed members of Gila are more closely related to Rhinichthys, Tiaroga, Agosia,

Lepidomeda, Meda, Plagopterus, Moapa, Algansea, Eremichthys, and Relictus than to G. atraria, their putative congener (Coburn and Cavender 1992: fig. 3). Actually, species of Gila in the subgenera

Temeculina and Gila are much more akin to members of Rhinichthys, Agosia, and several other genera than to G. atraria. The epithet Gila was initially assigned to forms which have since been subsumed within

G. robusta and G. elegans (Baird and Girard 1854a,b; see Jordan et al. 1930: 114 and Robins et al. 1991).

Since the Utah Chub belongs in a distinct taxonomic grouping that does not include those forms (Coburn and Cavender 1992), it should not be regarded as a member of the genus Gila.

Aforementioned morphologic evidence indicates that obesa and lineata were not synonymous, and not

Utah Chubs. Hence, Tigoma is not an appropriate generic epithet for the Utah Chub. Siboma (Girard 1857a:

26, c: 208) was the first generic cognomen assigned to any valid form of this species. Siboma atraria was taken at a Utah spring "near the desert" in 1853 (Girard 1857a,c, 1858b: 297-298, Cope and Yarrow 1876:

667-668, Tanner 1936). Snyder (1922) speculated that the species was collected at Fish Springs in southern

Tooele County based on the contradictory assumption that it was acquired in mid-May of 1854. In 1853, the unit of Captain J. W. Gunnison and Lieutenant E. G. Beckwith, which was searching for a potential railroad route to the Pacific Ocean, did not traverse Tooele County, but in October of that year, camped for a total of almost 21/2 days at Cedar Spring (near present-day Holden, Utah), situated in foothills at the margin of Black Desert (Beckwith 1854: 79-83). This is the most likely provenance of the single collected specimen of S. atraria. Siboma is an available name for the Utah Chub. Girard (1857a) solely assigned atraria to Siboma, and although he did not describe the genus, satisfied Indication 5 of Ride et al.'s (1985)

Article 16. Girard (1857c) described Siboma, after which he characterized crassicauda and then atraria. GREAT BASIN NATURALIST 12

Using software labeled Phylogenetic Analysis Using Parsimony (PAUP), Coburn and Cavender (1992) compared 48 cyprinid forms using 116 scale and osteological characteristics. The 48 compared forms did not all belong to the same taxonomic level. They included tribe, multi-generic, generic, subgeneric, and specific taxa according to current nomenclature (Robins et al. 1991). Analyses by Coburn and Cavender

(1992: fig. 3) show that at the highest level, the 12 forms in the Gila clade are divided into 2 nodes. One of these nodes comprises the Utah Chub and the Least Chub (lotichthys phlegethontis). The other embraces

10 other forms or taxonomic groups. The Least Chub, which may be a sister species of the Utah Chub, differed from the latter in only 4 of 116 characters (Coburn and Cavender 1992: 344-347, table 1). Also, the Utah Chub diverged from members of the subgenus Gila in only 4 characters. However, 3 of those 4 characters were osteological. In contrast, the Utah Chub and Least Chub differed in only 1 of 98 osteological characters but in 3 of 18 scale characters (Coburn and Cavender 1992, table 1). There is a tendency for cyprinid scale characters to be plastic and to exhibit convergence (repeated parallel trends) among disparate taxa, particularly within the subfamily Leuciscinae (Chu 1935: 74, 77), so scale structures may not be as taxonomically discriminating as osteological characters. Moreover, the Utah Chub and Least Chub appear to be uniquely similar in one important aspect. Examinations of six young Utah Chub specimens suggest that the posterior cranial fontanelle is not closed by parietal bones until relatively late in juvenile development

(Coburn and Cavender 1992: 346-347). In the Least Chub, this fontanelle persists. The retardation or absence of closure in the posterior cranial fontanelle is the trait on which the monophyly of these 2 species is based. More juvenile specimens of the Utah Chub need to be inspected to verify the delay in fontanelle closure. If further analyses confirm that the Utah Chub and Least Chub are sister species (see Coburn and

Cavender 1992: 344), the Least Chub would be called Siboma phlegethontis since lotichthys was not used as a generic epithet until much later (Jordan et al. 1930: 121) than Siboma (Girard 1857a: 26).

ED/HL .-,-% 14% in rhomaleus (TL 7.- 300 mm), 14.4 ± 1.8% in 1934-1935 specimens from Utah Lake

(X of TL = 280 mm; n = 16), 17% in atraria (in part) (TL = 283 mm; n = 9), 18% in squamata (TL 7--

200 mm) and atraria (TL = 178 mm [and = 508 mm]), 22% in cruoreus (TL = 152 mm), 23% (5.3 ± NATIVE PISCIFAUNA OF UTAH LAKE 13

0.3% of BL) in 1934 Utah Lake exemplars (g of TL 153 mm), .= 24% in pulverulentus (TL = 89 mm),

25% in bivittatus (TL = 76 mm) and domninus (TL = 51 mm), and 30% in timpanogensis (TL = 47 mm);

ED / interorbital width = 48.4% in 1934-1935 examples, 64.5% in 1934 specimens, and 83% in timpanogensis; ED/SL 2/3 in atrarius (in part; n = 9), = 1 in 1934 exemplars, and > 1 in timpanogensis

(Girard 1857a: 26, 1858: 297-298, Gill 1862b, 1876, Cope 1872b, 1876, Cope and Yarrow 1876: 654-655,

Jordan and Gilbert 1881, 1882: 195-196, 204, 234-235, 240-241, 886-887, Evermann 1893, Hayes 1935: derived from table 2, Carbine 1936: derived from table 5). There is a negative correlation between relative eye diameter (ED/BL) and BL (r = -0.624, df = 14, P < 0.01; derived from Hayes 1935).

Of the 6 cyprinids from Utah Lake and its affluents, the Utah Chub was the 2nd squattest (BD/BL =

27.2 ± 1.7%; n = 16) among 1934 specimens (Hayes 1935: derived from tables 2, 17-20, 22-23, 27-30,

35, 37). BD/BL = 23.9 ± 1.3% in 16 Utah Lake exemplars that were taken in 1934-1935, 25% in bivittatus, timpanogensis, and cruoreus, an average of 25.6% (n = 9) in atrarius from Jackson Hole,

Wyoming, 26.7% in pulverulentus, =26 or 28% in atrarius from Utah, an average of 27.2% in 16 Utah

Lake specimens collected in 1934, 28.6% in rhomaleus, and 33%, or 34% (30% of the distance from the tip of the snout to the end of the median caudal rays) in squamata (Gill 1862b, 1876, Cope 1872b, 1876,

Cope and Yarrow 1876: 654-655, Jordan and Gilbert 1881, 1882: 195-196, 204, 234-235, 240-241, 886-887,

Evermann 1893, Snyder 1922, Hayes 1935: table 2, Carbine 1936: table 5). In domninus, BD/BL = 20%

(Cope 1872b, Jordan and Gilbert 1882: 204), not even approaching that of its putative conspecifics. Only domninus and bivittatus had a horizontal gape (Cope 1872b, Jordan and Gilbert 1881, 1882: 195-196, 204,

240-241). Jordan and Evermann (1896a: 232-233) presumably synonymized domninus with the Utah Chub based on D, A, and 511, which, for the species typically = 8-9, 7-8, and 50-58, respectively (Sil = 59.7 ±

2.7 in 1934 examples) (Girard 1857c: 208, 1858b: 297-298, Gill 1862b, 1876, Cope 1872b, Cope 1876,

Jordan and Gilbert 1881, 1882, Snyder 1922, Hayes 1935, Carbine 1936). However, S. was lowest (= 9) in domninus; Sa = 10 in rhomaleus, 11 in cnioreus, 11.9 ± 0.5 in 1934-1935 exemplars, 11-13 in atraria,

12 in bivittatus, 13 in pulverulentus and timpanogensis, and 13.4 ± 0.7 in 1934 specimens. HL/BL = GREAT BASIN NATURALIST 14

<14 21.4% in bivittatus, 23.1% in domninus, 23.2 ± 0.8% in 1934 exemplars, slightly in atraria (in part),

'A in atraria and cruoreus, 26.0 ± 1.5% in 1934-1935 specimens, 26.7% in pulverulentus, 27.3% in timpanogensis, 28% in atraria, 28.6% in rhomaleus, and 30% or 35% in squamata; HL/TL = slightly > '4 in atraria and 28% in squamata (Girard 1857a: 26, 1858b, Gill 1862b, 1876, Cope 1872b, 1876, Cope and

Yarrow 1876, Jordan and Gilbert 1881, 1882: 195-196, 204, 234-235, 240-241, 886-887, Evermann 1893,

Snyder 1922, Hayes 1935, Carbine 1936). Because domninus possessed the combination of an inordinately slim body, small head, low S., and horizontal gape, it probably was not a Utah Chub. In bivittatus, P2 extended caudad only halfway to the anal fin (Cope 1872b). Sb = 5 in rhomaleus, 6 in timpanogensis, squamatus, cruoreus, and atraria, 6.5 + 0.6 in 1934-1935 exemplars, 8.2 ± 0.7 in 1934 examples, 9 in pulverulentus, but 11 in bivittatus (Girard 1857c, 1858b, Gill 1876, Cope 1872b, 1876, Cope and Yarrow

1876, Jordan and Gilbert 1881, 1882: 195-196, 234-235, 240-241, Evermann 1893, Hayes 1935, Carbine

1936). The uncharacteristically large SI, (11 versus a mode of 6), small head, and horizontal gape of bivittatus, as well as its uncharacteristically short P2 (Cope 1872b, Jordan and Gilbert 1882: 195-196) denote that it was not a Utah Chub either. The forms domninus and bivittatus will not be considered further.

In an addenda, Jordan and Gilbert (1882: 886-887) affirmed that their description of Squalius atrarius in the body of their monograph (p. 241) was based on juveniles of S. rhomaleus, and that Siboma atraria of Cope (1872b, 1876) (and presumably of Cope and Yarrow [1876: 667-668] who acquired 4 lots of Utah specimens) is synonymous with rhomaleus. Specimens that were labeled S. atraria by Cope (1872b, 1876)

(mass = 454-907 g) and Cope and Yarrow (1876) (mass 5 3175 g [.t = 455 g]; TL < 510 mm) were mature adults, and Squalius atrarius, the putative young rhomaleus, characterized as "a large species, used as food, and very destructive to young trout", was inexplicably larger (TL = 508 mm) than its supposed parent (TL = 305 mm), according to Jordan and Gilbert's (1882: 240-241, 886) recorded data.

Jordan and Gilbert (1882: 886) attested that it "is very doubtful" whether Girard's (1857a: 26, c: 208,

1858b: 297-298) Siboma atraria equals Cope's (1872b, 1876), and presumably Cope and Yarrow's (1876:

667-668) S. atraria. However, the described traits of the forms of Siboma atraria and Squalius atrarius NATIVE PISCIFAUNA OF UTAH LAKE 15

(Girard 1857a,c, 1858b, Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882: 241, 886-887) match quite favorably as Cope and Yarrow (1876) affirmed: "The specimens (of Siboma atraria) in our collection correspond well with Girard's description, except that in some instances we find the dorsal fin to have 10 rays; the ventrals 9" (likely due to polytypy; see above). Note that the expressions "9 + 1" and "8 + 1" that Girard (1858b) used to record D and A in the sole collected example of S. atraria (and also in T. obesa) obviously do not mean "n, or in some instances, n + 1", but presumably signify that the last ray was double. Only Snyder (1922) stated that D = 8 in Squalius atrarius. Jordan and Gilbert's (1882: 886-887) report that Sb = 7 in S. atrarius presumably reflects a misunderstanding of Girard's (1857c, 1858b) statement that "the lateral line in S. atraria runs along the seventh row of scales from the insertion of the ventrals (pelvic fins) upwards". Hence, Sb = 6 as Jordan and Gilbert (1882: 241) recorded.

The degree to which the decurved (Jordan and Gilbert 1882: 241) lateral line was complete was a function of age and size; it was incomplete in forms with TL 50 mm (Cope 1872b, 1876, Cope and

Yarrow 1876: 654-655, Jordan and Gilbert 1882: 196). Maxillae extended caudad as far as the rostral border of the eyes in cruoreus, rhomaleus, and squamata, almost that far in pulverulentus, but only as far caudad as the posterior margin of the nares in atraria (Girard 1857a: 25-26, 1858b: 297-298, Cope 1872b, Gill

1876, Jordan and Gilbert 1881, 1882: 234-235, 240-241, 886). Lobes of C (the caudal fin) were even in the

Utah Chub (Gill 1876, Jordan and Gilbert 1882: 234-235). The dorsal fin originated slightly craniad to P2

3 in timpanogensis, squamata, and some specimens of atraria, directly dorsad of 1 2's origin in rhomaleus and some examples of atraria, and slightly caudad to that origin in cruoreus and some specimens of atraria

(Girard 1858b, Gill 1862b, 1876, Cope 1876, Jordan and Gilbert 1881, 1882: 196, 234-235, 240-241,

Snyder 1922). Predorsal length = 50.6 ± 1.3% of BL in 16 specimens taken from Utah Lake in 1934

(Hayes 1935) and 54% of BL in atraria (Snyder 1922); the distance from the tip of the snout to P2 = 49.5

± 1.3% of BL in 1934 examples but 56% of BL in atraria. The length of P1 = 18.4 ± 1.0% of BL (n --

15) in 1934 specimens (Hayes 1935) and 20% of BL (16% of TL) in squamata (Gill 1876). P2's length ...--'

16.2% of BL (13% of TL) in squamata and 16.7 ± 1.2% (n = 16) in 1934 specimens. GREAT BASIN NATURALIST 16

A more detailed description is given for forms that have been taken from Utah Lake. As the Utah

Chub grows, its head becomes proportionately more depressed and its back more prominent, as exemplified in rhomaleus which had a broad, laterally concave head with a wide and flat interorbital span, a broad,

0 0 upturned snout, a rostrally protruding mandible, and a gape that formed angles of = 45 and .=--; 315 with the craniocaudal axis; it appeared dusky due to dense black speckling on a silvery background (Jordan and

Gilbert 1881, 1882: 240, Jordan 1891, Jordan and Evermann 1896a: 232-233). atraria (adult) appeared fuscous dorsally due to black stippling on an olive background; it was hoary ventrally (Girard 1858b:

297-298, Jordan and Gilbert 1882: 241). cruoreus (juvenile) had an anthracine dorsum, dusky sides, a red spot in axils of the pelvic and anal fins, an arched back, and a broad head that was rounded dorsally and was apparently straight or convex laterally; its lateral line coursed along a relatively low plane and premaxillae existed on the same horizontal plane as the bottom of the pupil (Jordan and Gilbert 1881, 1882: 234-235,

Jordan and Evermann 1896a). In the black-stippled and silvery-cheeked timpanogensis (juvenile), a narrow leaden line extending from the pterotic region to the base of C was superposed by brownish coloration and subimposed by yellowish (Cope 1876). Jordan and Evermann (1896a) remarked that scales of the Utah Chub are darkest along the edges where conspicuous lines may form. Rockwood (1873) commented that the modal mass of the Utah Chub .-,-; 450 g and the maximal mass < 1 kg. Jordan (1891) reported that 200 Utah Chubs netted in southern Utah Lake weighed < 575 g. Olson (1959: table 4) recorded that the maximal mass and length of Utah Chubs in Scofield Reservoir, Utah were 505 g and 308 mm, respectively.

Least Chub lotichthys phkgethontis (flaming small fish: Greek iota, small [pertaining to the smallest letter in the Greek alphabet]; ichthys, fish; phlegetho, to flame). If it is verified that the Least Chub is a congener of the Utah Chub (see preceding section), it would be known as Siboma phlegethontis. Synonyms are

Clinostomus phlegethontis (Cope 1876), Gila phlegethontis (Cope and Yarrow 1876: 657, Jordan 1878a:

424), Phoxinus phlegethontis (Jordan and Gilbert 1882: 244, 887, Jordan 1887b: 819), Hemitremia phlegethontis (Jordan 1891), Leuciscus phlegethontis (Jordan and Evermann 1896a: 243, b: 249), and lotichthys phlegethontis (Jordan et al. 1930: 121, Robins et al. 1991: 21). Its TL = 28-30 mm in specimens NATIVE PISCIFAUNA OF UTAH LAKE 17 from Juab County, Utah during 1976-1978 (Crawford 1979), 5 38 mm in pools about the mouth of Provo

River in 1889 (Jordan 1891), = 40.0 ± 4.6 mm in 8 examples that were collected from Provo River in 1934

(Hayes 1935: derived from table 29), 5 50 mm in other Provo River exemplars (Jordan and Gilbert 1882:

887), and < 65 mm in Utah (Holden et al. 1974). BL = 34 mm in Cope's (1876) specimens.

This species is the squattest of the 6 cyprinids from Utah Lake and its tributaries (Hayes 1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37). BD/BL = 28.6% in specimens from Utah's Beaver River (Cope

1876, Cope and Yarrow 1876: 657, Jordan and Gilbert 1882: 244) and 28.9 ± 1.6% in exemplars collected from Provo River in 1934 (Hayes 1935: derived from table 29). HL/BL = 23.5% (roughly estimated to be

14) in Beaver River specimens (Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882) and 25.2 ±

0.8% in Provo River examples (Hayes 1935). SL/BL = 6.82 + 0.4% in Provo River exemplars. ED/HL

= 31.4% (ED/BL = 7.9 ± 0.6%) in Provo River specimens whose BL = 40.0 ± 4.6 mm, and 36.4% in

Beaver River examples whose BL = 34 mm; ED / interorbital width ...--., 0.8 in the former and 1.2 in the latter (Cope 1876, Cope and Yarrow 1876, Hayes 1935). Relative ED (ED/BL) is negatively related to BL

(Cope 1876, Cope and Yarrow 1876; r = -0.875; df = 6, P < 0.005, Hayes 1935). The Least Chub has a blunt snout, short, oblique mouth, and maxillae that reach caudally to the rostral margin of the eyes

(Jordan and Gilbert 1882, Jordan 1891, Jordan and Evermann 1896a: 243, Hayes 1935). Jordan (1891),

Jordan and Evermann (1896a), and Hayes (1935) reported that the mandible projects rostrally beyond the upper jaw, but Cope (1876), Cope and Yarrow (1876), and Jordan and Gilbert (1882) indicated that the jaws were equal. Along a longitudinal row approximating the lateral line which has <7 pores, there were 36.0 ±

1.4 (mode of 35) scales in Provo River examples and 37 scales in Beaver River specimens (Cope 1876,

Jordan and Gilbert 1882, Jordan 1891, Jordan and Evermann 1896a, Hayes 1935). Cope (1876) and Cope and Yarrow (1876) noted that Sa + Sb = 11. According to Jordan and Evermann (1896a), Sa = 6 and Sb

= 4. Hayes (1935) reported that Sa = 6.0 ± 0.5 and Sb = 4.5 ± 0.5. A = 7.5 ± 0.8 in Provo River examples and 8 in Beaver River specimens; D = 7 in the latter exemplars, and had a mode of 8 in the former (Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882, Jordan 1891, Hayes 1935). The origin GREAT BASIN NATURALIST 18 of the dorsal fin is caudad to that of P2; predorsal length = 54.7% of BL in Beaver River specimens and

56.3 ± 2.6% of BL in Provo River exemplars; the mean distance from the tip of the snout to the origin of

P2 was 48.6 ± 1.2% in Provo River examples (Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert

1882, Jordan and Evermann 1896a, Hayes 1935). According to Cope (1876), P1 do not caudally come close to P2. Jordan's (1891) statement that P1 almost reach P2 is apparently in error. P2 extend to or beyond the origin of the anal fin (Cope 1876, Jordan 1891). Lengths of P1 and P2 were 17.8 ± 1.5% of BL and 15.9

± 1.7% of BL, respectively (Hayes 1935). Mean dorsal-fin height and basal length were 21.5 ± 1.0% and

12.4 ± 1.0% of BL, respectively; anal-fin depth and basal length were 18.6 + 1.3% and 11.8 ± 1.4% of

BL, respectively (Hayes 1935). This olivaceous or dark olive form has a dark vertebral streak, a broad dusky or plumbeous lateral streak, fuliginous stippling on its scales, and a faint spot on C (Jordan and Gilbert

1882, Jordan 1891, Jordan and Evermann 1896a, Hayes 1935, Holden et al. 1974). Cope (1876) declared that the dorsum is probably translucent in live individuals. The venter is golden or yellow, and may become washed with red, particularly in males during spring (Cope 1876, Jordan and Gilbert 1882, Jordan 1891,

Jordan and Evermann 1896, Hayes 1935, Holden et al. 1974). In males, fins are yellow or lemon amber and fin axils are red; females and juveniles may have silvery sides (Jordan 1891, Holden et al. 1974).

Longsnout Dace Rhinichthys transmontanus (trans-mountainous snout-fish: Greek rhinos, snout or nose; ichthys, fish; Latin transmontanus, trans-mountainous). The binomen of the Longnose Dace is Rhinichthys cataractae (Robins et al. 1991) whose type is Gobio cataractae in which "the muzzle is obtuse" (Cuvier and Valenciennes 1842: 315). Contrariwise, of the 6 cyprinids of Utah Lake and its tributaries, the

Longsnout Dace had proportionally by far the longest snout (11.0 ± 0.6% of BL) (Hayes 1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37). In G. cataractae, the rounded anal fin was longer ("wider") than the smaller dorsal fin, and BD/TL = only 12.5% (Cuvier and Valenciennes 1842). By comparison, in

Hayes' (1935: derived from table 30) 1934 examples of the Longsnout Dace from Provo River (n = 14), basal lengths of anal and dorsal fins were 10.9 + 0.7% and 12.7 ± 0.6% of BL, respectively; anal-fin depth and dorsal-fin height = 19.5 ± 0.8% and 21.1 ± 1.3% of BL, respectively. Although the Longsnout Dace NATIVE PISCIFAUNA OF UTAH LAKE 19 is the slenderest of the 6 cyprinids of Utah Lake and its tributaries (2 of BD/TL = 17.2%) (Hayes 1935: tables 2, 17-20, 22-23, 27-30, 35, 37), it is not nearly as slim as G. cataractae. BD/BL = 16.7% in

Rhinichthys luteus, 18.2% or 20% in R. maxillosus, 20% in R. cataractae, and 21.5 ± 1.3% in 1934 exemplars (Cope 1864, Garman 1880-1881, 1881, Jordan and Gilbert 1882: 207, Hayes 1935: table 30).

In G. cataractae, D = 9 and A = 8, but in Argyreus dulcis, R. maxillosus, R. transmontanus,

R. cataractae, and 1934 exemplars, D = 8 and A = 7 (Girard 1857c: 185-186, Cope 1864, Cope and

Yarrow 1876: 644, Jordan and Gilbert 1882: 207-208, Hayes 1935). Girard (1857c) indicated within the description of Argyreus nubilus that the ray counts for A. dulcis (D = 10 and A = 9) included 2 rudimentary rays; thus, D = 8 and A = 7. In several respects, Rhinichthys dulcis with HL/BL < 14, D =

9, A = 8, and a blunt snout overreaching a small mouth flanked by tiny or imperceptible barbels (Garman

1881, Jordan and Gilbert 1882: 885), resembled G. cataractae (Cuvier and Valenciennes 1842) more than

A. dulcis which had a pronounced snout, prominent maxillary barbels flanking a large mouth, and HL/BL

= '4 (Girard 1857a: 23, c, 1858b: 243-244). Jordan and Evermann's (1896a: 306-307) R. cataractae dulcis had a long, acute, and projecting muzzle, conspicuous barbels, and D, A, and SI, equal to 8, 7, and 63-70, respectively. None of these features were possessed by G. cataractae except that S11 = 70, the upper extreme of Su in R. c. dulcis. R. dulcis, with aforementioned traits and S1, = 80, does not fit either G. cataractae or its supposed synonym R. c. dulcis. G. cataractae, A. dulcis, and Rhinichthys spp. were all distinct.

Although Jordan (1880) inspected G. cataractae, comparisons herein show that there was not a sound basis for Jordan and Evermann's (1896a: 306-307, b: 261) synonymization of A. dulcis (Girard 1857a ,c, 1858b:

243-244), R. maxillosus (Cope 1864, 1872a,b, 1879, Jordan 1878a: 426, Garman 1881, Jordan and Gilbert

1882: 207), R. dulcis (Garman 1881, Jordan and Gilbert 1882: 885), R. transmontanus (= R. maxillosus of Cope and Yarrow [1876]) (Cope 1879, Garman 1881, Jordan and Gilbert 1882: 207-208), R. cataractae transmontanus (Jordan 1887b: 816), R. luteus (Garman 1880-1881, 1881), R. ocella (Garman 1881), R. cataractae (Jordan and Gilbert 1882: 207), and R. cataractae dulcis (Jordan 1887b: 815, Jordan et al. 1896) with that form. Later, Jordan et al. (1930: 139; see also Hayes [1935: 79] and Tanner [1936]) made GREAT BASIN NATURALIST 20

A. dulcis the type for western forms. Chute et al. (1948: 10) apparently lumped aforementioned forms and others within R. cataractae. Mayden et al. (1992: 838) regarded R. cataractae as polytypic.

Girard (1857c: 185-186; see also 1857a: 23) declared that examples of A. dulcis with a TL < 90 mm were taken in Sweetwater River, a Nebraskan tributary of Platte River, and that smaller specimens were acquired in Cottonwood Creek, Utah. Girard's (1858b: 243-244) more detailed account registers the collections of 9 adults and young in Sweetwater River and 1 juvenile in Cottonwood Creek. Snyder (1922) conjectured that the 4 taxa, A. dulcis, Cyprinella gunnisoni (Girard 1857c: 197) (= Notropis bubalinus

[Baird and Girard 1854c] based on Jordan's [1886b] statement that "The types of C. gunnisoni seem to be the young of N. bubalinus"; see also Jordan and Evermann [1896a: 273] and Jordan et al. [1930: 130] [any of the 12 examples of this small-headed (Girard 1858b: 267) form that remain in sound condition need to be reexamined and reidentified]), C. lugubris (Girard 1857c: 199) (likely = N. macrostomus; see Jordan and Gilbert [1882: 177], Jordan and Evermann [1896a: 274], and Jordan et al. [1930: 131]), and

C. ludibunda (Girard 1857c) (= Notropis ludibundus of Jordan and Evermann [1896a: 273]) and contrary to priority, N. stramineus of Robins et al. [1991: 77]; see also Jordan et al. [1930: 130]) were not collected at Cottonwood Creek, Utah as Girard (1857a: 23-24, c: 185-186, 197, 1858b: 243-244, 267, 271-272) noted but in an identically-named tributary of Arkansas River. Jordan and Evermann (1896a: 273) and Snyder

(1922) maintained that Notropis spp. do not occur in Utah or the Great Basin, but rather in the Mississippi

Basin. Snyder . (1922) did not account for the fact that A. dulcis and C. gunnisoni were acquired in 1854 when Lieutenant E. G. Becicwith's detachment was in Salt Lake Valley and further westward. Jordan et al.

(1930: 139) omitted the collection site of "Cottonwood Creek, Utah" in his synonymy of Rhinichthys (= A.) dulcis. C. lugubris and C. ludibunda as well as Bryttus (=Lepomis) humilis were garnered in 1853 (Girard

1858b: 21-22, 243-244, 271-272), so were most likely collected at the Cottonwood Creek mentioned by

Snyder (1922) which traversed the Santa Fe Trail 27 km from Lost Spring (Beckwith 1854: 17). These 3 taxa could have been taken from the Cottonwood Creek that parallels Utah Creek (W. A. Nichols [1857],

Map of the Department of New Mexico and Adjacent Territory; = Ute Creek [C. Roeser (1879), State of NATIVE PISCIFAUNA OF UTAH LAKE 21

Colorado (map), U.S. Department of the Interior, General Land Office]) in a northeasterly orientation a short distance to the west of Utah Creek where the J. W. Gunnison exploratory party camped for several days, but there is no record of any member of the unit going there (Beckwith 1854: 38-41). Noteworthily,

Girard (1857c: 199) reported that the provenances of C. lugubris and C. ludibundus were unknown.

Snyder's (1922) claim that Becicwith's (1854: 86-87) detachment camped on Salt Lake Valley's

Cottonwood Creek during 7 and 8 November 1853 is not true. When that party arrived in Salt Lake Valley on 8 November, men were mourning and livestock were exhausted (see below). Four days later, some 100 horses and mules were trailed to Cedar Valley to spend the rest of the late fall, and winter (Beckwith 1854:

5, 83, 86-87). Utah's Cottonwood Creek was not visited by Beckwith's (1854) unit any time in 1853. Of the 18 piscine taxa whose specimens Beckwith reportedly delivered to Smithsonian Institution, Girard

(1858b) noted that 10 were acquired in 1854, 9 paradoxically by a botanist, F. Creutzfeldt, who was killed in 1853. Girard (1858b) ascribed 4 of the first 5 to Beckwith, omitted the name of the collector of the 3rd of those species that were described by him, and recorded "Mr. Kreuzfeld" as the collector of the remaining

13 species, as if remembering while appraising the 6th taxon, of being informed that "Kreuzfeld" was in charge of biotic collections in Gunnison's unit (see Beckwith 1854: 2, 5). Girard (1857a) attributed the collections of all 18 to "Mr. Kreuzfeld" (a misspelling of the botanist's name). Snyder (1922) claimed that

Creutzfeldt and others were killed "just after entering" Salt Lake Valley on 8 November 1853. Actually, the massacre occurred on 26 October 1853 (Beckwith 1854: 79-83). The misinformation in Snyder (1922), the ignorance of Girard (1858b), and the vagueness about the provenances of multiple specimens that were putatively conspecifics of Utah Lake taxa, signal that the itinerary of Gunnison's exploratory party in central

Utah need to be elucidated (Beckwith 1854: 1-2, 78-87).

On a journey begun the previous 23 June, Gunnison's unit departed what is now Salina, Utah on 18

October 1853 and arrived in present-day Gunnison, 21.4 km to the north, early on 19 October. Captain

Gunnison and a military escort journeyed to Manti where he secured 2 Mormon guides who were brothers.

Meanwhile, the rest of the unit proceeded northward along Sevier River (roughly paralleling Utah Highway GREAT BASIN NATURALIST 22

28) until reaching, after 25.75 km, the proximity of today's Yuba State Park where the stream bottom narrowed so much that the party was forced to parallel Sevier River along adjacent hills until having traveled

28 km northward from Gunnison. After traveling 4 km downstream on 20 October, a road approximating today's Interstate 15 was encountered which extended between Salt Lake City and Los Angeles. On 21

October, the entire exploratory party continued southward and then southwestward on that road until reaching low foothills at Cedar Spring (or Cedar Springs) 16.4 km north of Fillmore in the vicinity of present-day Holden. On 22 October, Gunnison and an escort rode into Fillmore and arranged for agents of

Utah Territorial Governor Brigham Young to parley with Pah-Utah ( =Paiute or Piute) Indians to gain their assurances that their conduct toward the exploratory party would be peaceful. On 23 October, the unit headed northwest through Black Desert, the Mormon guides believing that water might exist in a mountain stream (perhaps on the isolated butte northeast of Pavant Butte) which they had seen flow in the spring.

However, this creek was dry and the party had to camp where there was neither grass nor water. After a grueling 24 October journey in sagebrush habitat contending with deep sand, the party reached Sevier River at the latitude of Delta (39°20'57" N), an estimated 40.9 km from Cedar Spring. Captain Gunnison, F.

Creutzfeldt, William Potter (Mormon guide), and 2 others with the escort of a corporal and 6 privates, set out on 25 October to surveil the surroundings of Sevier Lake. They followed Sevier River southwestward for an estimated 14.5 km to a point skirted by bird-inhabited sloughs where the stream begins to flow in a more westerly direction. After continuing > 3 km, Gunnison, concerned that Sevier River water further downstream might be too brackish for safe consumption, decided to camp at a cove (perhaps near the mouth of Dry Slough [Creek]) consisting of 2 sides flanked by grassland and willow groves, 1 side bordered by a high embankment, and 1 side constituting a passageway. While breakfasting there the next morning on 26

October, Gunnison's detachment was ambushed by Paiutes (the Gunnison Massacre National Monument exists 11 km east of Sevier River). The only ones who escaped slaughter were those who were able to quickly mount a horse and evade the bow- and rifle-wielding assailants after a long pursuit. Gunnison's body was pierced by 15 arrows and perhaps rifle balls as well. The 8 dead men included F. Creutzfeldt, botanist, and William Potter, Mormon guide. On 25 October, Becicwith's detachment had trudged 23.0 km NATIVE PISCIFAUNA OF UTAH LAKE 23 northeastward through sand, avoiding the largest dunes, and arriving east of present-day Lynndyl where the much-widened Sevier River branches to the south and to the east-northeast where the Sevier River gap lies

9.7 km distant between Gilson Mountains on the north and Canyon Mountains on the south. The next morning, Beckwith was awed by beholding through that gap, snow-clad Mount Nebo half-enveloped in clouds. He intended for his detachment to inspect the gap for potential wagon travel the following day. Later the same morning, Captain Morris and his subordinates had just left camp to determine the feasibility of establishing a route around the west side of Gilson Mountains and northward to Great Salt Lake, when approached by the corporal who had accompanied Gunnison southwestward, exclaiming that his comrades had been massacred. Morris and the remaining escort as well as the unit's physician, and G. G. Potter, brother of William, left on a rescue mission. At the murder scene, the forearms of several victims had been severed and detached, and their abdomen had been split to expose entrails from which "wolves" had already partaken. From east of Lynndyl, Becicwith's detachment, having been joined by 3 more survivors, proceeded

12.0 km southeastward on 26 October along a tortuous course to avoid deep sand, and immediately after stopping to set up camp, was united with the contingent of Captain Morris. On 27 October, the unit continued southward near the western margin of Canyon Mountains and then proceeded southeastward on the west side of Pavant Range, reaching Cedar Spring after journeying an estimated 45.7 km. Camp was moved to Pioneer Creek 4.8 km to the southeast on 28 October where grass for livestock was lusher. On

31 October, the exploratory party journeyed to the California road beside which it camped. Then on 1

November, it headed for Salt Lake City, arriving there on 8 November. En route, the unit camped along the wide and multi-channeled Timpanagos (= Provo) River on 4 November. On 12 November, the party's remaining horses and mules were trailed to Cedar Valley west of Utah Lake to graze and receive "several months' rest ... to recover from their weak and emaciated condition."

Jordan and Gilbert's (1882: 207) synonymization of their and Cope's (1864) (see also Garman 1881)

R. maxillosus was not warranted. S. + Sb = 25 in Cope's (1864) R. maxillosus but 19 in Jordan and

Gilbert's (1882). The largest contrast between Provo River specimens (Hayes 1935: table 30) and Jordan and Gilbert's (1882) R. maxillosus and R. cataractae was the sum of S. and Sb 26) in the former. Cope GREAT BASIN NATURALIST 24

(1864) noted that R. maxillosus had a "heavy" muzzle and a broad mouth and throat; he did not indicate that the snout was long and protuberant like that of Jordan and Gilbert's (1882) R. maxillosus and

R. cataractae, or Hayes' (1935: 80, table 30) Provo River specimens. Less importantly, Cope's (1864)

R. maxillosus was slimmer (BD/BL = 18.2%) than Jordan and Gilbert's (1882) R. maxillosus and

R. cataractae (BD/BL = 20%), or Hayes' (1935: table 30) Provo River examples (BD/BL = 21.4 ±

1.3%). ED/HL was slightly less in Cope's (1864)R. maxillosus (18.2%) than in Jordan and Gilbert's (1882)

R. maxillosus (19.0%) and R. cataractae (,--, 20%), or Hayes' (1935) Provo River exemplars (19.1%; ED/BL

= 5.0 ± 0.3%). TL was greater in Jordan and Gilbert's (1882) R. maxillosus and R. cataractae (127 mm) than in Cope's (1864) R. maxillosus (_ 75 mm) and Hayes' (1935) Provo River exemplars (X = 70.0 mm).

One anatomical agreement between Cope's (1864) R. maxillosus and Provo River specimens was the relatively caudad positioning of fins. Among Provo River examples, predorsal length = 55.1 ± 1.0% of BL, and the distance between the tip of the snout and the origin of P2 was 49.9 ± 1.3% of BL (Hayes 1935).

The dorsal fin of A. dulcis, by comparison, originated nearer the tip of the snout than the base of C (Girard

1857c: 185-186, 1858b: 243-244). Significantly, Sa = 11 in Jordan and Gilbert's (1882) R. maxillosus but

14.0 ± 0.7 in Provo River exemplars (Hayes 1935); Sb = 8 in R. maxillosus and R. cataractae, but 12.1

+ 0.5 in Provo River examples. Cope (1864) declared that fin size in R. maxillosus was small, and that P2 was particularly small, despite its caudad extension to the vent. Contrariwise, of the 6 cyprinids inhabiting

Utah Lake and its tributaries, the 2nd longest P1 ( = 21.3 ± 1.5% of BL) and the 3rd longest P2 ( = 16.2

± 0.8% of BL) were possessed by Hayes' (1935: tables 2, 17-20, 22-23, 27-30, 35, 37) R. dulcis. HL/BL in R. maxillosus, R. cataractae, and A. dulcis (Girard 1857a: 23, c, 1858b, Cope 1864, Jordan and Gilbert

1882) = '4, < the 26.3 ± 0.8% (n = 14) of Hayes' (1935: tables 2, 17-20, 22-23, 27-30, 35, 37) R. dulcis which had proportionally the greatest HL among the 6 cyprinids of Utah Lake and its tributaries (due partially to its protrusive snout). Barbels were markedly smaller in Provo River examples (Hayes 1935) than in Jordan and Gilbert's (1882) R. maxillosus and R. cataractae. The latter 2 forms differed from Hayes' NATIVE PISCIFAUNA OF UTAH LAKE 25

(1935) R. dulcis in so many ways that they must be regarded as distinct from the Longsnout Dace. Due to its prominent barbels, HL, and the position of its dorsal fin, A. dulcis is also a separate form.

R. luteus was chestnut with fuscous blotches and extensive black stippling, and had a dusky subnarial band; R. dulcis had a fulvous dorstun with light brown splotches, silvery venter, faint but dark lateral line, a fuliginous line passing from the snout through the eye, and a spot at the base of C (Garman 1880-1881,

1881, Jordan and Gilbert 1882: 885). R. ocella had a dusky dorsum with brown blotches and spots, a faint band passing from the snout and along the flank to C, and a yellow or orange venter in live individuals

(Garman 1881). In all 3 forms, D = 9, A = 8, maxillae reached caudad M of the distance to the rostral border of the eyes, the dorsal fin originated dorsad to the insertion (caudal basal attachment) of P2, and P2 extended caudad to the vent. Flanking a small, inferior mouth were small barbels in R. luteus and R. dulcis, but conspicuous barbels in R. ocella. The snout was protrusive in all 3 forms, but was also blunt in R. dulcis and R. ocella. HL/BL < IA in R. dulcis, 'A in R. luteus, and --,:: IA in R. ocella. S11 = 70 in R. ocella,

68-75 in R. luteus, but 80 in R. dulcis; in these 3 forms, S. = 11-12, 10-11, and 12-13, respectively. Sb =

10 in the first 2 forms but 12 in R. dulcis. ED/SL < 1/2 in R. luteus and R. ocella, but = V2 in R. dulcis.

Thus, these forms were likely different. Due to its small scales and head, R. dulcis was distinct from the

Longsnout Dace. Mainly because of its prominent barbels, R. ocella was also a separate form. Jordan (1891) created the subspecies R. dulcis luteus to embrace the exemplars of R. luteus which Garman (1880-1881) took in the proximity of Ogden, Utah, the "abundant" dace that Jordan encountered in Provo River near its mouth at Utah Lake, and in that lake's outlet, Jordan River, as well as some specimens that Jordan and

Gilbert (1881) ostensibly collected in Utah Lake and mislabeled Apocope vulnerata. Values of SI, in R. luteus

(68-75) and 1934 exemplars (67.3 ± 3.8) overlapped (Garman 1880-1881, 1881, Hayes 1935). However, it is ironic that Jordan (1891) regarded luteus to be a subspecies of R. dulcis for the purpose of comprising

Utah specimens having a high Sb ( = 14) when Sb in R. luteus = 10; also in R. luteus, Sa = 10-11, D = 8-9, and A = 7-8 (Garman 1880-1881, 1881). In Provo River examples, Sb = 11-14 (12.1 + 0.5), S. = 13-15

(14.0 ± 0.7), and D and A were invariably 8 and 7, respectively (Jordan 1891, Hayes 1935). BD/BL = GREAT BASIN NATURALIST 26

16.7% in R. luteus but 21.5 ± 1.3% in Provo River exemplars; ED / interorbital width (this width = 8.7

± 0.5% of BL in Provo River examples) = 57.7% in the latter but 2/s in the former (Garman 1880-1881,

1881, Hayes 1935). Its atypical features denote that R. luteus is not a Longsnout Dace.

Various parts of Jordan and Evermann's (1896a: 306-307) almost useless characterization of

R. cataractae dulcis (which nomenclaturally follows Jordan [1886b]) contradict the descriptions of various forms that are synonymized with that subspecies. For example, although Garman (1880-1881) and Jordan and Gilbert (1882: 885) indicated that R. luteus and R. dulcis had an obtuse muzzle, Jordan and Evermann

(1896a, 1900: plate 52) affirmed that R. cataractae dulcis had a long, projecting snout that was even

"sharper" than in other members of R. cataractae, and depicted the acute muzzle in a drawing.

According to Garman (1881), the principal difference between R. luteus and R. transmontanus was Sb which = 12-14 in the latter. Jordan (1887b: 816) declared that Sb in R. cataractae transmontanus was > that in any specimen of R. cataractae cataractae that he had ever seen. Sb in Provo River examples = 11-13

(12.1 ± 0.5) (Hayes 1935), but Jordan's (1891) only descriptive statement about R. dulcis luteus was that

Sb •=-,' a mode of 14 in Utah exemplars, but = 11-12 in R. dulcis dulcis. Since Sb = 10 in R. luteus from

Ogden (Garman 1880-1881, 1881), the examples with Sb = 14 necessarily came from Provo or Jordan River which, respectively, flow into and out of Utah Lake.

Of the putative forms of the Longsnout Dace, the only 1 to which specimens from Utah Lake and its affluents (Jordan 1891, Hayes 1935) anatomically approximate is R. transmontanus. There is 1 substantive difference between these forms. The dorsal fin of the latter was equidistant, presumably at its origin, between the tip of the muzzle and the base of C (Cope 1879). Contrariwise, predorsal length in Provo River examples 1,--, 55% of BL (Hayes 1935). Also, S. = 12-13 in R. transmontanus and 13-15 (14.0 ± 0.7) in

Provo River examples (Cope 1879, Cope and Yarrow 1876: 644, Garman 1881, Hayes 1935). The Provo

Longsnout Dace might be a distinct subspecies of R. transmontanus. Taxonomical studies are needed.

Western Dace Rhinichthys nubilus (dark snout-fish: Latin nubilus, dark or cloudy). Putative synonyms are

Argyreus nubilus, A. osculus, and A. notabilis of Girard (1857c: 186); A. nubilus of Girard (1858b: 244), NATIVE PISCIFAUNA OF UTAH LAKE 27

Ceratichthys nubilus, Apocope carringtonii, A. vulnerata, and Tigoma rhinichthyoides of Cope (1872b);

Alburnellus rhinichthyoides and Rhinichthys henshavii of Cope (1876); Apocope carringtonii, A. henshavii,

A. vulnerata, and A. oscula of Cope and Yarrow (1876: 645-647) who assigned C. nubilus of Cope (1872b) to A. henshavii, and placed part of the specimens of Cope's (1876) A. henshavii within A. oscula;

A. carringtoni, A. henshawi, A. vulnerata, A. oscula, A. notabilis, and A. nubilus of Jordan (1878a: 426);

Rhinichthys (Eritrema) rhinichthyoides of Garman (1880-1881), A. vulnerata (not all examples so labeled belong to this taxon [see Jordan 1891]) of Jordan and Gilbert (1881); A. carringtoni, A. nubila, A. vulnerata,

A. henshavii, and A. oscula of Jordan and Gilbert (1882: 209-211); Apocope vulnerata and Agosia novemradiata of Cope (1884); Agosia nubila and A. oscula of Jordan (1886b), Agosia novemradiata, A. carringtoni, A. nubila, and A. oscula of Jordan (1887b: 816), Agosia yarrowi (= Apocope oscula of Cope and Yarrow [1876: 647] but not Argyreus osculus of Girard [1857c: 186]) and A. nubila of Jordan (1891);

A. yarrowi, A. nubila, and A. nubila carringtonii of Jordan and Evermann (1896a: 309-312, b: 262); and

Apocope nubila, A. carringtonii, and A. yarrowi of Jordan et al. (1930: 140-141). Hayes (1935: 86-87) considered C. nubilus, but not Argyreus nubilus, to be a synonym of Apocope carringtonii which also comprised carringtonii, vulnerata, rhinichthyoides, and henshavii. Bailey et al. (1960: 16) and Robins et al.

(1991: 24) regarded R. nubilus as a synonym of R. osculus. Jordan and Evermann (1896a) provided an inaccurate and incomplete summary of descriptive data. The variable (Jordan 1896a, Robins et al. 1991: 78)

Western Dace is presumed to be polytypic (see Mayden et al. 1992: 839).

Several phenotypic measurements of the Western Dace and Longsnout Dace overlap; muzzle length is the most discriminating. In the former taxon, the lateral line is incomplete in small specimens, maxillae extend caudally ,-- 2/3 the distance to the rostral border of the eyes, and an inferior mouth is flanked by filiform barbels that are minute or imperceptible (Cope 1872b, Cope and Yarrow 1876: 646-647, Garman

1880-1881, Jordan and Gilbert 1882: 209-210, Jordan and Evermann 1896a: 311-312). HL/TL = 20% in nubilus and henshavii, 20.5% in 1934 exemplars, and 21% in oscula; HL/BL 23% in vulnerata and T. rhinichthyoides, 24% in nubilus, carringtonii, and henshavii, 25% in R. rhinichthyoides and novemradiata, GREAT BASIN NATURALIST 28 and 25.5 ± 0.9% in 1934 examples (Girard 1857c: 186, 1858b: 244, Cope 1872b, 1876, 1884, Cope and

Yarrow 1876: 645-647, Garman 1880-1881, Jordan and Gilbert 1882: 209-211, Jordan 1887b: 816, Hayes

1935). ED / interorbital width = 2/3 in oscula, carringtonii, vulnerata, henshavii, and T. rhinichthyoides, and ---, 72% (interorbital breadth = 9.2 + 0.5% of BL) in 1934 exemplars (Cope 1872b, Cope and Yarrow

1876, Jordan and Gilbert 1882, Jordan 1887b, Hayes 1935). ED/HL = 20% in nubilus whose TL

100 mm and in vulnerata whose TL..-- 50 mm, ',.--- 22% in oscula whose TL < 75 mm, novemradiata whose TL = 107 mm, and R. rhinichthyoides, .-=-, 23% in henshavii, --= 24% in carringtonii whose TL =

42 mm (and 100 mm), = 25% in vulnerata whose TL = 64 mm, > 1,4 in vulnerata and in

T. rhinichthyoides whose TL --.-; 50 mm, and averaged 25.7% in 1934 specimens (ED/BL = 6.6 ± 0.6%) whose TL 'A,- 58 mm (BL = 46.4 ± 8.0 mm) (Cope 1872b, 1876, 1884, Cope and Yarrow 1876, Garman

1880-1881, Jordan and Gilbert 1882, Hayes 1935). Relative ED is negatively correlated with BL (see above).

BD/TL.--.., 13% in oscula and 17-18% in henshavii; BD/BL .-=-z 19% in henshavii and T. rhinichthyoides,

20% in carringtonii and R. rhinichthyoides, 19-22% in vulnerata, 20-21% in novemradiata, 24% in nubilus, and 25.9 ± 2.0% (n = 10) in 1934 examples (Cope 1872b, 1876, 1884, Cope and Yarrow 1876, Garman

1880-1881, Jordan and Gilbert 1882, Jordan 1887b, Hayes 1935). The large BD in 1934 Provo River exemplars may be an artifact of small sample size. In 51 specimens that Hayes (1935: table 34-37) collected from other Utah waterways, BD/BL = 23.6 ± 1.2%.

HL/BL = 23.3% in 7: rhinichthyoides of Cope (1872b) and 'A in R. rhinichthyoides of Garman

(1880-1881). ED/SL = 2/3 in the latter but 1 in the former; ED / interorbital breadth = 2/3 in the former.

Based on examples of Hayes' (1935: tables 35 and 37) carringtonii from Provo River, relative eye diameter

(ED/BL) is negatively correlated with BL (r = -0.943, df = 8, P < 0.001) so the difference between the

ED/HL of 22.2% in R. rhinichthyoides and > 'A in T. rhinichthyoides is not taxonomically meaningful.

Su = 60 in carringtonii, nubilus, and novemradiata, 63 in Apocope oscula, 64.3 ± 1.7 (n = 10) in

1934 examples from Provo River (in other Utah waterways, minimal Sp = 60 in 1934), 62-67 in henshavii,

65 in vulnerata, 65-67 in forms of rhinichthyoides, and 90 in Argyreus osculus (Cope 1872b, 1876, 1884, NATIVE PISCIFAUNA OF UTAH LAKE 29

Cope and Yarrow 1876: 645-647, Garman 1880-1881, Jordan and Gilbert 1882: 209-211, Jordan 1887b:

816, 1891, Hayes 1935: derived from tables 34-37). Sll :-- 70-72 in vulnerata (Jordan and Gilbert 1881); part of the specimens so labeled were judged by Jordan (1891) to be the Longsnout Dace (as were some of

E. D. Cope's Apocope specimens) whose mean Su and Sb are higher (see Hayes 1935). Sb = 9.8 ± 1.1 in

Provo River specimens, 10 in oscula, 10-12 in henshavii, 11 in carringtonii and novemradiata, and 10-14 in vulnerata and forms of rhinichthyoides; Sa = 10 in oscula and carringtonii, 11 in novemradiata, 12-13 in forms of rhinichthyoides, 12-15 in vulnerata, 13.6 ± 0.5 in 1934 exemplars, and 14-16 in henshavii

(Cope 1872b, 1876, 1884, Cope and Yarrow 1876, Garman 1880-1881, Jordan and Gilbert 1882, Jordan

1887b, Hayes 1935: table 37). Noteworthily, Sb in Hayes' (1935: tables 34-37) examples of carringtonii from Utah waterways other than Provo River (12.7 ± 1.0 [n = 51]) was > Sb in exemplars from that river.

Sa values were 10 or 11 in 3 forms but 13 in 61 examples of the Western Dace taken from various Utah sites in 1934 (Hayes 1935). Jordan (1891) manifested that Argyreus osculus had tinier scales than Apocope oscula (and other putative synonyms), and thus, is a distinct form that will not be considered further.

The predorsal lengths of 55.3% of BL in novemradiata and 56.8 ± 1.5% of BL in 1934 examples do not differ much; the distance from the tip of the snout to P2 = 51.8% of BL in the former and 51.8 ± 1.8% of BL in the latter (Cope 1884, Hayes 1935). Unlike most putative forms of the Western Dace, D = 9 in novemradiata; however, D also = 9 in variant 1 of henshavii and in 2 of 10 exemplars collected in 1934, and = 9-10 in R. rhinichthyoides (A = a mode of 7 in 1934 specimens and = 7 in other forms) (Girard

1857c: 186, 1858b: 244, Cope 1872b, Cope 1876, Cope and Yarrow 1876: 645-647, Garman 1880-1881,

Jordan and Gilbert 1882: 209-211, Jordan 1887b: 816, Hayes 1935). ED/HL = 22.2% in novemradiata

(Cope 1884) and R. rhinichthyoides (Garman 1880-1881) but averaged 25.7% in 1934 exemplars (Hayes

1935). ED/SL = 213 in novemradiata and R. rhinichthyoides, almost % (SL/BL = 8.8 ± 0.7% of BL) in

1934 examples from Provo River, < 1 in henshavii (includes Ceratichthys nubilus of Cope [1872b]), and

= 1 in vulnerata and T. rhinichthyoides (Cope 1872b, Cope and Yarrow 1876, Garman 1880-1881, Jordan and Gilbert 1882, Jordan 1887b, Hayes 1935). Confirming that novemradiata had a long snout, Cope (1884) GREAT BASIN NATURALIST 30 commented, "The head is rather elongate, especially the muzzle, which projects a little beyond the mouth" and is longer that the snout of henshavii. The depth of the "rather deep" (Jordan 1887b) caudal peduncle in novemradiata (10.6% of BL) (Cope 1884) was < that in all but 1 of the 6 cyprinids of Utah Lake and its tributaries; the caudal peduncle in 1934 examples of the Western Dace from Provo River (n = 10) was the longest (23.8 ± 1.5% of BL) and deepest (12.7 ± 0.8% of BL) among the cyprinids of Utah Lake and its affluents (Hayes 1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37). The combination of the slender caudal peduncle, long snout, D, and small ratios of ED/HL and ED/SL signify that novemradiata is distinct from the Western Dace. In R. rhinichthyoides, those ratios were also small and D's value (9-10) was high (Garman 1880-1881); Garman (1880-1881), although segregating the tally of the 2 anterior rudimentary rays of this form's dorsal fin, might have counted the branches of a double posterior ray as 2 rays. Nonetheless, R. rhinichthyoides should not be regarded as a Western Dace at least until all sound remaining examples of the 70 specimens of this form that Cope (1884) collected, are reexamined.

The dorsal fin of the Western Dace originated slightly caudad to the point of origin of P2; the latter approached or reached the anal fin caudally, but P1 did not extend caudad to P2 (Cope 1872b, 1876, Cope and Yarrow 1876: 645-647, Jordan and Gilbert 1882: 209-211). Hayes (1935) confirmed that predorsal length (56.8 ± 1.5% of BL) is > prepelvic length (51.8 ± 1.8% of BL). Girard (1857c: 186) attested that both dorsal and anal fins are "well developed". Jordan and Evermann (1896a: 311-312) corroborated that their heights equal. Hayes (1935: derived from tables 2, 17-20, 22-23, 27-30, 35, 37) showed that among the 6 cyprinids of Utah Lake and its tributaries, the difference in the heights of dorsal and anal fins is next to the least in the Western Dace (1.4% of BL). This dace's dorsal-fin height = 22.6 ± 1.3% of BL, and its anal-fin depth = 21.2 ± 1.2% of BL. Ventral fins can be reddish, crimson, or pinkish, at least in the breeding season (Cope 1872b, 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882, Hayes 1935).

Information on coloration is provided for forms that have been collected in Utah Lake or Provo River.

The Western Dace is gray with scattered dark spots or blotches on the body as well as on dorsal and anal fins and dorsal part of the head (Jordan and Gilbert 1882: 209-210, Jordan and Evermann 1896a: 311-312, NATIVE PISCIFAUNA OF UTAH LAKE 31

Hayes 1935). A broad, fuscous lateral band extended from the muzzle to C in vulnerata and straight back from the eye as far caudad as the anal fin in "oscula"; a dusky longitudinal filament existed on vulnerata's thoracic region (anterior venter), and in "oscula", a narrow black stripe continued rostrad from the anterior margin of the eye to and around the tip of the muzzle, passing ventrad to the nares (Cope 1876, Cope and

Yarrow 1876: 646-647, Jordan and Gilbert 1882: 210-211, Jordan and Evermann 1896a, Hayes 1935: 87).

According to Cope and Yarrow (1876: 647) and Jordan and Gilbert (1882: 211), "oscula" possessed an ivory dorstun, gamboge venter, dusky vertebral line that extended at least as far caudad as the dorsal fin's origin, rosy lower lip, and rosy lower jaw in males. vulnerata had an olive or olivaceous brown dorstun with dusky spots that were concentrated to form a blackish vertebral line, and possessed a yellow venter, head that was blackish at least dorsally, and red chin (in males); henshavii was generally white or pallid but had red lips, a fuliginous wash on the head, and sparse dusky splotches on the caudal peduncle and sides (Cope

1876, Cope and Yarrow 1876: 645-647, Jordan and Gilbert 1882: 210). Hayes (1935) declared that the axils of the dorsal and anal fins in 1934 specimens were pink during the breeding season. In henshavii and

"oscula" , ventral fins were reddish or crimson, at least in their axils (Cope 1876, Cope and Yarrow 1876,

Jordan and Gilbert 1882: 210-211). Juveniles of henshavii had dusky transverse streaks (Jordan and Gilbert

1882: 210). Hayes (1935) noted that specimens from murky waters are lighter than those from clear water.

Bonneville Redside Richardsonius hydrophlox (Richardson's flame-water: Richardsonius refers to Sir John

Richardson, M. S., F.R.S., F.L.S., surgeon and naturalist on the first British expedition under the command of Captain Sir John Franklin to a region of North America between 54-67° N latitude, and the person who described Cyprinus (Abramis) balteatus based on a specimen from Columbia River that was sent to him by

Dr. Gairdner, surgeon of Hudson Bay Company's post at Fort Vancouver, Oregon Territory along that river;

Greek hudro, water; phlox, flame). This form was apparently considered by Chute et al. (1948) and definitely considered by Robins et al. (1991: 24) to be included in the species Richardsonius balteatus (=

Cyprinus [Abramis] balteatus) (Richardson 1836: 301-302, Girard 1857c: 202, 1858b: 278-279). However, in balteatus, the dorsal fin, but particularly the slightly concave anal fin whose length exceeded its depth, GREAT BASIN NATURALIST 32 were proportionately longer and lower than those fins in the Bonneville Redside. Anal-fin basal length and depth in C. (Abranu:s) balteatus (Richardson 1836) were 25.0% and 16.4% of BL, respectively; dorsal-fin basal length and height were 14.7% and 19.0% of BL, respectively. By comparison, those dimensions in examples of the Bonneville Redside from Provo River = 14.9 ± 1.9% of BL (n = 61), 19.7 ± 1.3% of

BL (n = 60), 12.6 ± 0.9% of BL (n = 61), and 20.9 ± 1.3% of BL (n = 61), respectively (Hayes 1935: derived from tables 17-20). Richardson (1836) intimated that he may have counted an initial rudimentary ray in the anal and dorsal fins of C. (Abramis) balteatus; his sums were A = 19-22 and D = 11. Girard

(1858b) indicated that in 2 examples of R. balteatus measuring 127 mm and 152 mm, A = 19 and 21 (+

2 rudimentary rays in both), respectively, and D = 11 (+ 2 rudimentary rays; the datum for D in specimen 2 was missing). In the Bonneville Redside, A = 10-12 and D = 8-9 according to Cope (1872b,

1876), Cope and Yarrow (1876: 657-660), and Evermann (1893); Jordan (1891) stated that maximal A =

13. The basal length of the anal fin was > HL (18.8% of TL) in R. balteatus (Girard 1858b) but anal-fin length (g = 11.9% of TL) was much shorter than HL = 18.7% of TL) in the Bonneville Redside (Hayes

1935). Hence, balteatus the Bonneville Redside. Not unexpectedly, Mayden et al. (1992: 839) considered

R. balteatus to be polytypic. Putative synonyms of the Bonneville Redside are Clinostomus hydrophlox and

C. montanus (Cope 1872b), C. hydrophlox and C. taenia (Cope 1876), Gila montana, G. hydrophlox, and

G. taenia (Cope and Yarrow 1876: 657-660, Jordan 1878a: 424), Squalius taenia and S. montana (Jordan and Gilbert 1881), S. hydrophlox, S. taenia, and S. montanus (Jordan and Gilbert 1882: 233-234), Phoxinus hydrophlox, P. taenia, and P. montanus (Jordan 1887b: 818), Leuciscus montanus (Jordan 1891),

L. hydrophlox (Evermann 1893, Jordan and Evermann 1896a: 238-239, b: 248), Cheonda montana and

C. hydrophlox (Jordan et al. 1930: 119), and Richardsonius hydrophlox (Hayes 1935, Tanner 1936). La

Rivers and Trelease (1952) called this taxon R. balteatus hydrophlox.

Although taenia was seldom seen in Utah Lake, specimens of it were taken in Provo River near its mouth at Provo, Utah; hydrophlox was acquired from Utah Lake where it was abundant and montanus was

collected from Provo River at Provo, Utah where it was "very abundant" (Cope 1872b, 1876, Cope and NATIVE PISCIFAUNA OF UTAH LAKE 33

Yarrow 1876: 657-660). In montanus and taenia, the jaws were equal; the mandible of hydrophlox projected rostrally beyond the upper jaw (Cope 1872b, Cope and Yarrow 1876, Jordan and Gilbert 1882: 233-234).

Jordan and Evermann (1896: 238) noted that the head of hydrophlox might be distinctly narrower than that of the other 2 forms which Jordan (1891) and Evermann (1893) declared were identical. In the Bonneville

Redside, Su = 56-58 and A = 10-13 (mode of 10 or 11); hydrophlox, with S. = 15, Sb = 7, and D = 8, may be distinct at some taxonomic level from the other 2 forms in which S. = 11-12, Sb = 5-6, and D =

9 (Cope 1872b, 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882, Jordan 1891, Jordan and Evermann

1896a). According to Rockwood (1873), the mass of the Bonneville Redside = 110-230 g. In Hayes' (1935)

61 examples of R. hydrophlox (comprising the 3 aforementioned forms) from Provo River, mean scale and ray sums were Su: 56.1 ± 2.2, S.: 11.6 ± 0.8, Sb: 6.3 ± 0.8, D: 8.9 ± 0.4, and A: 10.6 ± 1.0.

Jordan and Evermann (1896a: 238) did not provide an adequate or accurate summary of the characteristics of the Bonneville Redside. Its HL and SL were proportionally < those in Rhinichthys spp.

(see above). HL/BL = 21% in hydrophlox, .--,-- 23% (HL/TL = 19%) in taenia, 23.3 ± 1.1% in 1934 specimens, and 25% in montanus; SL/BL = 6.4 ± 0.7% (n = 61) in 1934 examples (Cope 1872b, 1876,

Cope and Yarrow 1876: 657-660, Jordan and Gilbert 1882: 233-234, Hayes 1935: derived from tables

17-20). The Bonneville Redside has a short mouth with an oblique gape, anal fin that is longer than the dorsal fin, and lateral line, sometimes incomplete, that is deflexed at a place dorsad to the midpoint between

P1 and P2; P2 do not extend caudally to the anal fin (Cope 1872b, 1876, Cope and Yarrow 1876: 658-659,

Jordan and Gilbert 1882, Jordan and Evermann 1896a, Hayes 1935: 57). Evermann (1893) reported that the dorsal fin existed 7-- midway between the tip of the snout and the base of C, but Hayes (1935) indicated that the dorsal fin originated further caudad (predorsal length = 55.0 ± 2.4%). BD/HL was almost 1 in montanus, = 1 in taenia, and averaged 1.0 in 1934 exemplars; BD/BL z-,-, 22% in hydrophlox, .-- 23% or

25% in taenia, = 24.1 ± 1.3% in the 1934 specimens, and = 25% or almost 25% in montanus (Cope

1872b, 1876, Cope and Yarrow 1876: 657-660, Jordan and Gilbert 1882, Hayes 1935: tables 17-20).

Maxillae reached caudally to the rostral border of the eyes in hydrophlox and taenia but beyond that point GREAT BASIN NATURALIST 34 in montanus (Cope 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882). ED/HL = 20% in hydrophlox whose TL ,--- 150 mm, = 29% in montanus whose TL = 90 mm, 29.4% (ED/BL = 6.8 ± 0.6%) in 1934 examples whose TL averaged = 83 mm (BL = 66.8 ± 8.9 mm), and = 31% in taenia whose associated

TL = 73 mm (Cope 1872b, 1876, Cope and Yarrow 1876, Jordan and Gilbert 1882, Hayes 1935). Ocular diameter / interorbital breadth (this width = 8.4 ± 0.5% of BL in 1934 exemplars) = 2/3 in hydrophlox,

0.81 in 1934 Provo River examples, and 1 in montanus and taenia (Cope 1872b, 1876, Cope and Yarrow

1876, Hayes 1935). Based on Provo River specimens that were taken in 1934 by Hayes (1935), there was an inverse correlation between relative eye diameter (ED/BL) and BL (r = -0.622, df = 59, P < 0.001).

Cheeks of the Bonneville Redside are golden or crimson (apparently in males), and silvery or silvery ocher (evidently in females), and may bear a bright silver or golden crescent (Cope 1872b, Cope and Yarrow

1876: 658, Jordan and Gilbert 1882: 233, Hayes 1935). The suboculus may be crimson or turquoise, and a golden supercilium may pass from the snout to the gill opening (Cope 1872b, Jordan 1891, Jordan and

Evermann 1896a: 238). Some specimens from Spanish Fork River had iridescent purplish green sides (Hayes

1935: 57). In multiple examples, the dorsum was largely greenish silver or silvery (iridescent) green, but dusky on the upper back (Cope and Yarrow 1876: 658-659, Jordan and Gilbert 1882: 234, Jordan and

Evermann 1896a, and Hayes 1935). A silvery band existing above the middle Vs of the body might be subimposed by a bright saffron stripe which in turn is subimposed by a black filament (Cope and Yarrow

1876); alternatively, a fuliginous or anthracine lateral band might be sandwiched between 2 silvery stripes

(Jordan and Gilbert 1882, Jordan and Evermann 1896a, Hayes 1935). This lateral vitta and/or the region below it, were also characterized as being bright orange in females and red or crimson in males, or as being bright scarlet or crimson only in breeding males (Cope and Yarrow 1876: 657-658, Jordan and Gilbert 1882,

Jordan and Evermann 1896a). According to Jordan (1891) and Jordan and Evermann (1896a), the red lateral band of males did not continue further caudad than the origin of the anal fin. Hayes (1935) noted that the venter of breeding males was fiery scarlet between P1 and the anal fin. The olive upper dorsum of hydrophlox was subimposed by a longitudinal dusky border even with the top of the eye, which was NATIVE PISCIFAUNA OF UTAH LAKE 35 subimposed by a crimson band that in turn was subimposed by a dark vitta coursing immediately dorsad to the lateral line and extending from the epiclavicular region to C; the venter was crimson anteriorly and silvery posteriorly, or entirely silvery in females (Cope 1872b, Cope and Yarrow 1876: 658, Jordan and

Gilbert 1882: 233). Cope (1872b) and Cope and Yarrow (1876: 657-658) stated that montanus was colored similarly, except that the venter was everywhere crimson, but Jordan and Gilbert (1882: 234) affirmed that the sides were red only in males. In taenia, the venter was white, the iris was black, and the much larger female lacked bright lines (Cope and Yarrow 1876: 658-660); in some examples, sides were silvery as far dorsad as the lateral line which was superposed by a fuliginous vitta extending from the tip of the snout to

C, that in turn was superposed by a white line which was superposed by a dusky dorsum (Cope 1876).

Individuals from alkaline or muddy waters may be pallid (Jordan and Gilbert 1896a, Hayes 1935).

Family Catostomidae

Exceptional Sucker Catostomus generosus (exceptional low-mouth: Greek kato, low; stoma, mouth; Latin generosus, superior or exceptional). Putative synonyms are Acomus generosus (Girard 1857a: 21, 1858b:

221-222), Catostomus (Acomus) generosus (Girard 1857c: 174), C. generosus (taken at Utah Lake) (Cope

1872b), Minonuis platyrhynchus, M. jarrovii, and C. ?generosus (all 3 collected about Provo, Utah) (Cope

1876), and Pantosteus platyrhynchus and P. jarrovii (Cope and Yarrow 1876: 673-675). Neither

C. generosus nor the 1.1-kg C. ? generosus was described. To be able to confirm that they resemble

A. generosus more than M. platyrhynchus or M. jarrovii, sound examples of these 2 generosus forms must persist, allowing their comparison with descriptions of the 2 latter forms, and any remaining intact exemplars of them. After comparing specimens, Jordan (1878a: 416, b: 183-184) deemed A. generosus (= C. (Acomus) generosus) equal to P. jarrovii, but P. platyrhynchus a separate species. Jordan (1878b) (see also Jordan and

Gilbert [1882: 1231) stated that P. generosus and P. platyrhynchus were very similar but added, "at present,

I consider (them) distinct." Later, Jordan (1886b) proclaimed, "The type of Catostomus generosus seems to me unquestionably identical with the type of" P. platyrhynchus and "with P. jarrovii". "The peculiarities of P. platyrhynchus seem to be due to its shriveled condition." Actually 3 specimens (apparently syntypes) GREAT BASIN NATURALIST 36 of A. generosus (Girard 1858b) and several syntypes of M. plaiyrhynchus (Cope 1876) had been collected.

Jordan (1887b: 805) also lumped all 3 form within P. generosus. Jordan (1891) and Jordan and Evermann

(1896a: 170-171) regarded Cope's (1876) M. jarrovii and M. platyrhynchus as synonyms of A. generosus

(Girard 1857a,c, 1858b) but Cope and Yarrow's (1876: 674-675) P. jarrovii as a synonym of Catostomus plebeius which had been collected at Mimbres River near the border of USA and Mexico (Baird and Girard

1856). Likewise, Jordan (1887a) and Jordan and Evermann (1896b: 238, 1902: 45-46) did not consider platyrhynchus or jarrovii to be full species. Jordan and Evermann (1896a: 170) gave an inaccurate and incomplete synopsis of P. generosus (which included M. platyrhynchus and M. jarrovii). Snyder (1922) speculated that A. generosus had been taken from the Rio Grande Basin, and that A. generosus might equal

Catostomus plebeius. In accordance with Snyder's (1922) paper, Jordan et al. (1930: 104) replaced

P. generosus with P. platyrhynchus as the nominal form, and noted that M. jarrovii from Provo, Utah was a questionable synonym of P. platyrhynchus because "the type" (actually there were 2 syntypes) of

M. jarrovii (Cope 1876) had been lost. Contrary to priority but in accordance with Jordan et al. (1930),

Moore (1968) referenced the taxon using the contemporary binomen (Robins et al. 1991), C. platyrhynchus.

Those fishes for which collection localities were recorded that Beckwith reportedly delivered to

Smithsonian Institution, were reportedly caught in Utah's Cottonwood Creek, a Utah spring "near the desert", Colorado's Utah Creek, and a sluice in Arkansas River near Fort Makee (Girard 1857a: 21-26,

1858b: 21-22, 221-222, 236-237, 243-244, 267, 271-272, 277-278, 297-298, 320-321). Beckwith possibly conveyed to Smithsonian Institution the exemplars of the 3 taxa that were taken in the Arkansas River sluice but Pimephales maculosus was not collected by the Gunnison/Beckwith party in 1853 because it did not pass through Fort Makee (Beckwith 1854). Furthermore, F. Creutzfeldt did not collect any of the 3 forms. All other exemplars from tributaries of Arkansas River were taken by Dr. C. B. Kennerly, H. B. Mollhausen, and Dr. George G. Shtunard serving under the command of Lieutenant A. W. Whipple (Girard 1857a: 23,

1857c: 166, 170, 178-179, 182, 189-191, 193, 198, 1858b: 4-5, 14-17, 21-22, 25-26, 28-29, 96-98,

208-209, 211-212, 231, 234-237, 249-251, 256, 259-263, 266-268, 270, 275-276). Gunnison's unit surveyed NATIVE PISCIFAUNA OF UTAH LAKE 37

0 a land swath largely lying along 37.5-40 N latitude. After Gunnison's death and a winter sojourn in Salt

Lake Valley, Beckwith directed the party's search for a railroad route along = 41° N latitude (Beckwith

1854: 1, 86-87, Girard 1857a,c). The course of Utah's Cottonwood Creek lies at ,--,v 41°10-15' N latitude.

In an article rife with inaccuracies (e.g., noting that specimens of Gila elegans and Algansea obesa were of unknown origin when in actuality, G. elegans had been collected at 5 different known sites including

Grand River, Utah, and A. obesa had been taken in Humboldt River [Baird and Girard 1854a, Girard 1857a:

22, 25, c: 183, 205, 1858b: 239, 286-287]), Snyder (1922; "do" in his table means "ditto") indicated that

6 piscine species that had been taken in 1853 or 1854 at Cottonwood Creek, "an affluent of the great Salt

Lake of Utah" (Girard 1857a: 21, 23-24, c: 174, 185-186, 197, 199, 1858b: 21-22, 221-222, 243-244, 267,

271-272), were actually from the Mississippi River Basin. Four of these 6 taxa are treated above in the section on the Longsnout Dace. The other 2 were A. generosus , and Bryttus (= Lepomis) humilis (see Jordan and Evermann [1896a: 10041). Of the 10 examples of B. humilis that Girard (1858b: 21-22) recorded (see also Girard 1857a: 21, 1858a), only 1 was obtained from Cottonwood Creek. Apparently, Snyder's (1922) skepticism about the origin of fishes that had been taken in Cottonwood Creek, Utah was triggered by Jordan and Evermann's (1896a: 273) comment that the recorded collection site of Cyprinella gunnisoni (likely =

Notropis bubalinus) must be in error because Notropis spp. do not occur in Utah (refer to the section on the

Longsnout Dace). Snyder (1922) reasoned that if any specimens (e.g., those of Notropis spp.) which were reported to have been taken from Cottonwood Creek, Utah were not actually collected there, none of them were. In accordance with the publication and comments of Snyder (1922) who was a colleague and former student of Jordan's at Stanford University, Jordan et al. (1930: 130-131), without a sound evidentiary basis, proclaimed acquisitions of C. gunnisoni from "Cottonwood Creek of the Rio Grande, Colo.", C. ludibunda from a "locality now known to be Cottonwood Creek, Colo.", and C. lugubris from "Cottonwood Creek,

Colo." Snyder (1922) declared that Beckwith's detachment was without a naturalist on 8 November 1853 when it supposedly camped at Cottonwood Creek, Utah so "it was quite probable that no collecting was done." However, Beckwith's (1854) party did not visit that stream in 1853, and furthermore, collected GREAT BASIN NATURALIST 38

C. gunnisoni in 1854 (Girard 1858b: 267) when the unit was in Salt Lake Valley and further westward.

Snyder (1922) believed (probably wrongly) that Siboma atraria was taken in Utah during 13-16 May 1854.

If that were the case, the same collector could have also acquired A. generosus and Argyreus dulcis at

Cottonwood Creek. Snyder (1922) related that J. A. Snyder, the unit's assistant topographer, had "collected some specimens." He could also have done so at Cottonwood Creek. Near the end of his report dated 31

January 1854, Beckwith (1854: 98) signified that a replacement(s) had been designated to describe biological collections, a work that had been delayed because of Creutzfeldt's death. Girard (1858b: 221-222) stated, and Girard (1857c: 174) implied that A. generosus was collected in Cottonwood Creek, Utah by Beckwith.

Snyder (1922) asserted that Cottonwood Creek, Utah "is... as is now evident, an impossible locality for the species which (Girard) assigned to it", and proclaimed that A. generosus was not really taken in

Bonneville Basin because other species reportedly collected there actually inhabit Mississippi Valley.

Ironically, Snyder (1915) had earlier attested, "It is quite within reason to presume that (Notolepdomyzon

= Acomus) generosus is represented in the upper Columbia Basin." Even though Girard (1857a: 21, c: 174,

1858b: 221-222) indicated that A. generosus was collected in Cottonwood Creek, Utah, Snyder (1922) did not ascribe a stream with that name as its provenance, but conjectured, "It now seems probable that the fishes called A. generosus were collected in the Rio Grande basin, perhaps in Utah Creek" where Salar virginalis was taken (see the section on the Bonneville Cutthroat Trout). Snyder's (1922) justification for this opinion was the resemblance that he noticed between Catostomus plebeius from that basin and A. generosus.

It is ironic that Snyder felt constrained to speculate on a non-Utah collection site for A. generosus when

C. generosus, C. ? generosus, M. platyrhynchus, and M. jarrovii were all collected about Utah Lake (Cope

1872b, 1876). Jordan et al. (1930: 104) quoted Snyder as stating that A. generosus had been taken not in

Utah's Cottonwood Creek but the one "no doubt from Colorado." In Costilla County of southern Colorado, there is a Cottonwood Creek that parallels Ute (= Utah) Creek (C. Roeser [1879], State of Colorado [map],

U.S. Department of the Interior, General Land Office) in a northeasterly orientation a short distance to the west of it. However, Girard (1858b: 320-321) was fundamentally accurate in stating that Utah Creek was NATIVE PISCIFAUNA OF UTAH LAKE 39 a tributary of Rio Grande del Norte; if collections had been made in a neighboring stream, it seems likely that associated specimen labels would have noted that that waterway was also a tributary of the same river.

The adjacent Utah and Cottonwood Creeks are tributaries of Trinchera Creek (or River) which flows into northern Rio Grande River (see also U.S. Department of the Interior, Geological Survey [1973], State of

Colorado [map]). Subsequent to Snyder's articles (1915, 1922) and remarks, Jordan (1930: 104) placed

A. generosus, C. plebeius, and 2 other species in a distinct genus, Notolepidomyzon.

In Utah Lake forms, scales are smaller cranially than caudally, the head bears conspicuous mucous tubes, HL/TL = 17.4-21.4%, Su = 79-87, S. = 14-17, and ST, = 11-14 (Cope 1876, Cope and Yarrow

1876: 673-675, Jordan and Gilbert 1882: 123, Snyder 1922). The distended lower lip possesses 4-5 rows of papillae; the upper lip has 2 (Jordan and Gilbert 1882). Labial folds are pronounced and tuberculate; a notch exists where the posterior fold (which is deeply incised caudally) joins the anterior fold (Cope 1876,

Cope and Yarrow 1876). Labial folds were discoid-infundibular in platyrhynchus, and narrow rostrally in jarrovii. In platyrhynchus, the transverse and abruptly angular labial commissure was covered with a cartilaginous sheath, C was openly emarginate, P, extended caudad halfway to P2, and the latter did not reach the vent (Cope 1876, Cope and Yarrow 1876: 673-674). The light brown jarrovii had dusky spots and blotches, a faint abdominal stripe, red fins, and a rostrally-convex and sharp-edged labial commissural sheath

(Cope 1876, Cope and Yarrow 1876: 674-675). The examples of platyrhynchus collected about Provo, Utah had a dusky dorsum and pinldsh yellow venter (Cope 1876, Cope and Yarrow 1876: 673-674).

Although some of their traits are similar, BD/TL = 13.5% in platyrhynchus but 17.4% in M. jarrovii;

D = 11 in the former but 9 or sometimes 10 in the latter; the muzzle was depressed and very protuberant in the former but tapered dorsally and only slightly protrusive in the latter; ED / interorbital width = 60% in the former but 50% in the latter; predorsal length = 47.0% in the former and 50.4% in the latter; in platyrhynchus, the orbit's superior border was indented (Cope 1876, Cope and Yarrow 1876: 673-675).

These 2 forms seem to be distinct but the differences between the 107-mm M. jarrovii and the 168-mm

, platyrhynchus might be age-related. In A. generosus, predorsal length :.--. 50% and P2 originated ventrad to GREAT BASIN NATURALIST ao

1 the caudal /3 of the dorsal fin; in platyrhynchus and M. jarrovii, the distance from the snout's tip to P2 =

55.0% and 55.7%, respectively (Girard 1857a,c, 1858b, Cope 1876, Cope and Yarrow 1876, Jordan and

Gilbert 1882: 123, Snyder 1915, 1922). generosus (BD/TL = 20-22.2%) was squatter than M. jarrovii and particularly, than platyrhynchus (see above); generosus lacked a frontoparietal fontanelle whereas platyrhynchus had a long, narrow one. Thus, generosus and pknyrhynchus are likely separate. Mayden et al. (1992) regarded the Mountain Sucker as polytypic. Utah exemplars must be compared and re-identified.

Mountain Sucker Catostomus platyrhynchus (broad-snouted low-mouth: Latin platy, broad; Greek rhunkhos, snout or bill). This species is largely treated in the preceding section. Catostomus generosus (Cope

1872b) and C. ? generosus (Cope 1876), which were not described, and Minomus jarrovii (Cope 1876; see above) could possibly be Mountain Suckers as well. It is crucial to be able to identify the larval Mountain

Sucker so that it can be differentiated from the endangered (Lowe et al. 1990) June Sucker during recovery-program surveys. Analyses of supposed larval June Sucker habitats that were undertaken in June of 1982-1984 were in vain; larval suckers that were collected from lower Provo River in those years (not

1982, 1983, and 1986 as indicated) and sent to the Colorado State University Larval Fish Laboratory for identification were all determined to be Mountain Suckers (Radant et al. 1987: 8, 10). Moreover, those larvae that were garnered from Provo River in 1985 and placed in an isolated body of water for rearing were later identified as Mountain Suckers (Radant et al. 1987: 10). Shirley (1983) described the ontogeny of the putative June Sucker via illustrations and textual data. However, almost all of the collected metalarvae (final larval stage) and juveniles, as well as some of the younger larvae upon which his analysis was based, were actually Mountain Suckers (Snyder and Muth 1988: 3). Snyder and Muth's (1988: 26-27, figs. 8-15, 37-43) identification guide demonstrates that the amount of melanistic pigment on the upper back and mid-ventral line of larval June and Utah Suckers (Catostomus ardens) is generally less than that in larval Mountain

Suckers. In the first 2 larval stages, the number of myomeres between the tip of the snout and the vent was

36 ± 1 (n = 22) in the Mountain Sucker but 38 + 1 (n = 13) in the June Sucker; predorsal length in the

3rd and 4th (last) larval stages were 48 ± 1% (n = 10) and 49 ± 1% (n = 12) of BL, respectively, in the NATIVE PISCIFAUNA OF UTAH LAKE 41

June Sucker but 50 ± 1% (n = 11) and 51 ± 1% (n = 9) of BL, respectively, in the Mountain Sucker; dorsal-fin lengths in those larval stages were 13 ± 1% (n = 9) and 17 ± 1% (n = 9) of BL, respectively,

in examples of the Mountain Sucker, 15 ± 1% (n = 5) and 19 ± 1% (n = 12) of BL, respectively, in the

Utah Sucker, and 17 ± 1% (n = 5) and 22 ± 2% (n = 12) of BL, respectively in the June Sucker (Snyder

and Muth 1988: tables 6, 11, 16). The sums of myomeres existing between the tip of the snout and the

origin of the dorsal finfold in the 1st and 2nd larval stages were 12 ± 1 (n = 10) and 12 ± 0 (n = 5),

respectively, in the Utah Sucker, but 15 ± 1 in both stages (n = 13) of the June Sucker (Snyder and Muth

1988: tables 6 and 11). In juveniles of the Mountain Sucker and Utah Sucker, the mouth and mandibles

migrate caudally as the young fish matures such that the mouth becomes inferior; contrariwise, the mouth

and mandibles of the June Sucker move little (Snyder and Muth 1988: figs. 16-17, 30-31, 44-45).

Unknown catostomid. This fish from Utah Lake resembled C. ardens but had a wider lower lip with 8-10

rows of smaller, coarse, irregular papillae (Jordan and Gilbert 1881). The dorsal lobe of the large C was

14 the length of the head. This form's very long P1 were almost as long as its head, and P2 extended caudad

to the vent (Jordan and Gilbert 1881). The breast was almost scaleless. Unlike other Utah Like catostomids,

the free posterior margins of the gill membranes formed a broad fold across the narrow isthmus, as in

Coitus. Jordan and Gilbert (1881) opined that if the structure of the isthmus persists during continued growth

and development, this form probably ought to be considered to belong to a distinct genus.

Utah Sucker Catostomus ardens (burning low-mouthed one: Latin ardens, burning or flaming; in reference

to a bright rosy lateral band that appears on males at least during the breeding season). In their original

description of C. ardens, Jordan and Gilbert (1881) suspected that a specimen which Henry Crecy Yarrow

and H. W. Henshaw had collected at Utah Lake in July 1872, and which Cope and Yarrow (1876: 679) had

labeled Catostomus guzmaniense (sp.), might also belong to this species. Cope and Yarrow (1876) assigned

that specimen to Catostomus (Aconms) guzmaniensis (Girard 1857c: 173) whose syntypes had been collected

in Janos River, a tributary of Guzman Like in Chihuahua Province, Mexico (Cope and Yarrow 1876), far

removed from Bonneville Basin. Subsequently, guzmaniensis was subsumed within Pantosteus GREAT BASIN NATURALIST 42

(= Notolepidomyzon = Catostomus) plebeius (Jordan 1891: 20, Jordan et al. 1930: 104, Robins etal. 1991).

Since 1881, the Utah Lake specimen (guzmaniense) has not been mentioned. The characteristics of the Utah

Sucker are compared with those of other Utah Lake catostomids in the next section on the Webug Sucker.

Webug Sucker Chasmistes fecundus (fecund yawner: Greek chasmao, to yawn or gape; Latin fecundus, fecund). Four lots of specimens of C. fecundus that had been acquired in Utah Lake by Henry Crdcy Yarrow and H. W. Henshaw in 1872 were described and compared with Catostomus (Acomus) generosus by Cope and Yarrow (1876: 678-679). Upon examination of some of these fecundus specimens, Jordan (1878a, b:

149-151) concluded that they belonged to an undescribed genus which he differentiated from Catostomus and named Chasmistes. Then in the addendum to the latter treatise, Jordan (1878b: 219-220) indicated that after reexamining Cope and Yarrow's (1876: 678-679, plate 32) textual description and drawing of fecundus, his notes on those authors' "typical" specimens of fecundus, and the specimens on which his description of the genus Chasmistes was based, Jordan determined that the specimens of Chasmistes belonged to a species distinct from fecundus which he named liorus. Hence, Jordan's (1878a, b: 149-151) description of fecundus actually refers to liorus. Liter, Jordan (1882: 129) provided the first substantive characterization offecundus based on monophyletic specimens. Although Jordan (1878b: 219-220) had earlier furnished the initial information on the Webug Sucker after it had been differentiated from the June Sucker, most of that characterization was simply a reiteration of Cope and Yarrow's (1876: 678-679) description. After analyzing a specimen at the U.S. National Museum that Jordan (1878b: 219-220) considered to be typical offecundus, he affirmed that fecundus belongs to the genus Catostomus rather than Chasmistes. Ironically, the mouth of that particular specimen was in poor condition so he was unable to evaluate its lips. Jordan (1878b: 149-150) had previously noted that the mouth and lips are of "singular" importance in differentiating Chasmistes from

Catostomus. Jordan (1878b: 219-220) assigned 2 lots of specimens from the U.S. National Museum that had been collected at Utah Lake by Yarrow and Henshaw, to fecundus. Subsequently, the Webug Sucker was known as Catostomus fecundus (Jordan and Gilbert 1881, 1882: 129, Jordan 1891, Jordan and Evermann

1896a: 180-181) until it was again placed in the genus Chasmistes (Jordan and Evermann 1896b: 240, 1898b: NATIVE PISCIFAUNA OF UTAH LAKE 43

2794, Jordan et al. 1930: 108). Tanner (1936) believed that there was but 1 species of sucker in Utah Lake which he referenced as Catostomus fecundus. However, his view later changed. Lowder (1951: 1), one of

Tanner's students, observed 6783 suckers for his thesis project. Finally, after inspecting ,----- 6700 of these fish, Lowder (1951) concluded that they actually belonged to 3 species, and took measurements of the remaining individuals. Another student under Tanner's advisement (Stubbs 1966: 6, 10) inspected > 1000 catostomids, identifying a few Webug and Utah Suckers as well as numerous June Suckers. In taxonomically categorizing mature adult ( 3 yr of age) catostomid specimens, Stubbs (1966: 1-4, 11) referred to their descriptions by Cope and Yarrow (1876: 678-679), Jordan and Gilbert (1881), Jordan and Evermann (1896a:

179-183), and perhaps Jordan (1891); he did not allude to the generic or specific treatments of these forms by Jordan (1878b: 149-151, 219-220) and Jordan and Gilbert (1882: 128-129, 131-132). Stubbs' (1966: iii,

11, tables 1-3) thesis advisor verified his identifications, and ultimately 25 specimens representing the 3 year-round catostomids of Utah Like were selected for detailed osteological examinations. Stubbs (1966:

52) referred to the Webug Sucker as Catostomus fecundus as his advisor had done (Tanner 1936), but concluded that the Webug Sucker is much more closely related to the June Sucker than to the Utah Sucker, and that the Webug and June Suckers have a common immediate ancestor, thus corroborating the position of Jordan and Evermann (1896b, 1898b) and Jordan et al. (1930) that fecundus belongs in the genus

Chasmistes. Based on Miller and Smith's (1981: 11) contention that fecundus is not a distinct taxon, Robins et al. (1991) did not include it in their checklist of North American piscine species. Miller and Smith (1981), who did not make reference to Jordan and Evermann (1896b, 1898b) and Jordan et al. (1930), alluded to the Webug Sucker as Catostomus fecundus in accord with Jordan and Evermann (1896a).

Note that the description of Chasmistes (Jordan 1878b: 149-150) leads to the inference that it was based on both the Webug and June Suckers even though this description was restated by Jordan and Gilbert (1882:

131), and to a large extent by Jordan and Evermann (1896a: 182), during the period that the genus was considered to be monotypically represented by the June Sucker. Some generic traits that are detailed in that description such as "anal fin high (deep), reaching caudal", "pectorals rather long, not quite reaching GREAT BASIN NATURALIST 44 ventrals (pelvic fins)", and "eyes ... high up, rather posterior" conform to characterizations of the Webug

Sucker rather than the June Sucker (see below). The ascription of an evenly lobed tail to Chasmistes by the genus description is not warranted by the features of either sucker.

It was Stubbs' (1966: 8, 51) view that the shape of the head and snout, and the position of the mouth are the external features that best differentiate the year-round catostomids of Utah Lake. With reference to the anterior dorsum's contour, Stubbs (1966: 29, fig. 42) noted that the dorsal surface of the Webug Sucker is "moderately decurved" rostrad from its back which, according to Jordan (1881), is "a little elevated".

Jordan and Gilbert (1881, 1882: 129), Jordan (1891), and Jordan and Evermann (1896a: 180-181) reported that the dorsal surface of the Webug Sucker is regularly decurved from the nape to the base of the premaxillary spines which project rostrally on roughly a horizontal plane (Cope and Yarrow 1876: plate 32) to form a markedly protuberant snout. Predorsally, the dorsal profile of the June Sucker is almost horizontal, according to Stubbs (1966: 29). Jordan (1878b: 149-150), Jordan and Gilbert (1881, 1882: 131), and Jordan and Evermann (1896a: 182) indicated that the head of the June Sucker is dorsally concave or depressed, low at the nape, and highest at the tip of the prominent upturned snout and the caudal extremity of the head. The interorbital span is markedly convex in the Webug Sucker, but very broad and nearly flat in the June Sucker

(Jordan and Gilbert 1881, 1882: 132, Jordan and Evermann 1896a: 183). In the Utah Sucker, the premwdllae are scarcely elevated above the level of the mandibles (Jordan 1881). In the Webug Sucker, the caudal peduncle is long and thin, and the lower lobe of the slender, moderately forked tail is longer and wider than the upper; the June Sucker's lower caudal lobe is also the longer one, but its caudal peduncle is rather stout (Jordan 1878b: 149-150, Jordan and Gilbert 1881, 1882: 129, 131-132; Jordan and Evermann

1896a: 180-181, 183). In the Utah Sucker, the short and broad tail is roughly even (Jordan and Gilbert 1881,

1882: 128-129, Jordan 1891, Jordan and Evermann 1896a: 179-180). From the characteristics of Utah Lake catostomids, Stubbs (1966: 44-45, plate 5) deduced that the Utah Sucker is the most elongate and fusiform with the narrowest back and shoulders, and that the June Sucker has the broadest back and bulkiest shoulders. Jordan and Gilbert's (1881) account confirms that the June Sucker is heavy at the shoulders. NATIVE PISCIFAUNA OF UTAH LAKE 45

Although the publications of Jordan and Gilbert (1881, 1882: 128-129) and Jordan and Evermann (1896a:

179-180) corroborate that the Utah Sucker has a rather elongate, subfusiform body with a dorsally convex and subconic head, they also indicate that the slightly elevated back of this species is wide.

The mucous-channeled head of the June Sucker is widest dorsally, and has sides that may be directed somewhat ventromediad according to the description of Chasmistes (Jordan 1878b: 149-150, Jordan and

Gilbert 1882: 131). In the Utah Sucker, the head is also broad, at least dorsally (Jordan 1881, Jordan and

Evermann 1896a: 179-180). In contrast, Stubbs (1966: 44-45, plate 5) stated that the Utah Sucker has the smallest (cuneate or subconic) head. Recorded cephalic lengths of Utah Lake catostomids are on a crude average, smallest for the Webug Sucker at 20% (Cope and Yarrow 1876: 678-679; part of the specimens were this species), 23% (Stubbs 1966: derived from table 7), or 25% (Jordan and Gilbert 1882: 129, Jordan and Evermann 1896a: 180-181) of BL. HL/BL = 20% (Cope and Yarrow 1876: 678-679; part of the specimens were this species), 23% (Stubbs 1966), or 27% (the mode) in the June Sucker (Jordan and Gilbert

1882: 132, Jordan and Evermann 1896a: 183, Miller and Smith 1981: the mean derived from table 1), and

20% (Stubbs 1966), 25% (Jordan 1891), or 27% in the Utah Sucker (Jordan 1881, Jordan and Evermann

1896a: 179-180). The front of the Utah Sucker's small (ED/HL ..-- 14%) eye exists about midway in the head; in the Webug Sucker, the eye is positioned slightly caudad to the middle of the head (Jordan and

Gilbert 1881, 1882: 128-129, Jordan and Evermann 1896a: 180-181). In the latter, ED/HL ,=:: 14% (Jordan and Gilbert 1881) or 17% (Cope and Yarrow 1876: 678-679; part of the specimens were this species).

Jordan (1881) affirmed that rostrocaudally, the eye of the June Sucker lies exactly at the midpoint of the side of the head; ED/HL ,--:-. 11-13% (Miller and Smith 1981: table 1) or 14-17% (Cope and Yarrow 1876: 678-

679 [part of the specimens were this species], Jordan and Gilbert 1881, Jordan and Gilbert 1882: 132, Jordan and Evermann 1896a: 183). In the Utah Sucker, the thick, very wide, and pendent upper lip comprises 4-5

(Jordan 1891), 4-8 (Jordan and Evermann 1896a: 179-180), or ..--: 8 (Jordan and Gilbert 1881, 1882: 128-129) rows of coarse, irregular papillae that may be largest in the second and third rows from the lingual side; the lower lip is very broad, deeply incised medially across its entire width, and composed of 2 narrowly spaced GREAT BASIN NATURALIST 46 lobes. The Webug Sucker's pendent, rather smooth, and somewhat protrusive upper lip contains ,--- 4 rows of large, coarse papillae, and a wide medial gash separates the 2 lobes of the lower lip on which ,-- 6 rows of papillae exist; lips of the June Sucker are smooth and nonpapillose or nearly so (Jordan 1878b: 149-150,

1891, Jordan and Gilbert 1881, 1882: 129, 131, Jordan and Evermann 1896a: 180-182, Holden et al. 1974,

Shirley 1983). The rather protrusive upper lip of the June Sucker is very thin; each of the widely separated lobes of the medially incised lower lip lies laterad to a mandible, is weakly folded with slightly uneven plicae, and narrows rostrally to become a medial rim (Jordan 1878b: 149-150, Jordan and Gilbert 1881,

1882: 131, Jordan and Evermann 1896a: 182, Shirley 1983). Holden et al. (1974) claimed that the June

Sucker's upper lip is concealed by an overhanging snout. Its robust mandibles, which 7-, 35-45% of HL and

0 0 become proportionately larger with age, form angles of ,...- 45 and ,---- 315 with the craniocaudal axis (Jordan

1878b: 149-150, Jordan and Gilbert 1881, 1882: 131; Jordan and Evermann 1896a: 182). In the Webug

Sucker, large mandibles form angles of 60° and 300° with that axis (Jordan and Gilbert 1881, 1882: 129,

Jordan and Evermann 1896a: 180-181), or are almost as oblique as those of the June Sucker (Jordan 1891).

Smaller mandibles bordering the Utah Sucker's inferior mouth are horizontal (perpendicular to the body axis) or only slightly oblique, and measure <¼ of HL (Jordan and Gilbert 1881, 1882: 128-129, Jordan 1891,

Jordan and Evermann 1896a: 179-180). The very large mouth of the June Sucker and the smaller mouth of the Webug Sucker (Jordan 1878b: 149-150, 219, 1891, Jordan and Gilbert 1882: 131, Jordan and Evermann

1896a: 182, Sigler and Sigler 1987: 239-241, Page and Burr 1991: 167-168) ) are terminal or subtenninal.

Stubbs (1966: 44-45) related that the Utah and Webug Suckers have relatively high fins and that the

June Sucker has proportionately the smallest fins. In partial corroboration, Jordan and Gilbert (1881) affirmed that the anal fin is deep in the Webug Sucker, and very deep and long in the Utah Sucker, reaching the base of C caudally. Jordan and Gilbert (1882: 128-129) and Jordan and Evermann (1896a: 179-180) characterized the anal fin of the Utah Sucker as long rather than deep. Jordan and Gilbert (1881, 1882: 129,

132) and Jordan and Evermann (1896a: 180-181) confirm that the Webug Sucker's anal fin is deep, and the

June Sucker's ventral fins are small. Jordan (1891) mistakenly indicated that the June Sucker's fins are large. NATIVE PISCIFAUNA OF UTAH LAKE 47

Jordan (1891) stated that the Webug Sucker's ventral fins are long; indeed, the relatively long P1 certainly extend > 1/2 of the distance between their origin and that of P2 (Jordan and Gilbert 1881, 1882: 129, Jordan and Evermann 1896a: 180-181). The Utah Sucker's P1 are short and broad, and its squat P, do not reach the vent caudally (Jordan and Gilbert 1881, 1882: 128-129, Jordan 1891, Jordan and Evermann 1896a:

179-180). In the Webug Sucker, the dorsal fin's anterior ray is about twice the height of the posterior ray, and longer (Jordan and Gilbert 1881, 1882: 129; Jordan and Evermann 1896a: 180-181) or almost as long

(Jordan 1891) as that fin's basal length. In the June Sucker, the anterior ray is about twice the height of the posterior ray and = 83% (Jordan 1891) or 100% (Jordan and Gilbert 1881, 1882: 132, Jordan and

Evermann 1896a: 183) of the basal length. The height of the anterior ray of the Utah Sucker's dorsal fin is only 1.5 times that of the posterior ray and 75-100% of that fin's basal length (Jordan and Gilbert 1882:

128-129, Jordan 1891, Jordan and Evermann 1896a: 179-180). Jordan and Gilbert (1881) noted that the free margin of the June Sucker's dorsal fin is almost straight, but Jordan (1891) related that it is slightly concave.

According to Jordan and Gilbert (1881, 1882: 129), the Webug Sucker is dusky dorsally due to dense dark stippling, and pallid or silvery ventrally. Jordan (1891) noted that it is rather pale. In the June Sucker, the olive dorsum is so profusely and darkly stippled or mottled that it appears dusky; the venter is pallid or white, and the fins are pale except at their fuliginous tips (Jordan and Gilbert 1881, 1882: 132, Jordan 1891,

Jordan and Evermann 1896a: 183). The Utah Sucker has a dusky dorsum that is blotched with black, sides that are vaguely spotted, a white venter, and fins or ventral fins that are mottled with fuliginous (Jordan

1881, 1891, Jordan and Gilbert 1882: 128-129; Jordan and Evermann 1896a: 179-180). Rockwood (1873) commented that the general coloration of this species varies from crimson to dark green. The Utah Sucker is darker than the Webug Sucker (Jordan 1891) and the June Sucker (Rockwood 1873), but at least anteriorly, the male (Jordan and Gilbert 1881, 1882: 128-129; Jordan and Evermann 1896a: 179-180) or breeding male (Sigler and Sigler 1987) possesses a bright rosy lateral band. The adult Utah Sucker of Utah

Lake had a TL of 46 cm (Jordan and Gilbert 1881, 1882: 128-129, Jordan and Evermann 1896a:

179-189), a modal mass of 0.7 kg, and a maximal mass 2.3 kg (Rockwood 1873) ( < 1 kg according GREAT BASIN NATURALIST 48 to Jordan [1891]). In the 1800s, the June Sucker's TL typically ..--' 46 cm (Jordan and Gilbert 1881, 1882:

132, Jordan and Evermann 1896a: 183) and its maximal mass = 1.4 kg (Jordan 1891). Because there has been little recruitment of young into the June Sucker population in recent years, this taxon's average size has been continually increasing (Gutermuth and Lentsch 1993: 2-3, Gutermuth et al. 1993: 2, 14-15, table 3). The mean mass of 126 suckers (of which 81% were June Suckers) that were netted in Utah Lake in 1978-1979 was 1.6 kg (Radant and Sakaguchi 1981: 141). Mean mass = 2.4 kg among 35 spawning June

Suckers in 1991 (Gutermuth and Lentsch 1993: table 1, omitting the captive specimen), and 2.3 kg among

57 spawning June Suckers in 1992 (Gutermuth et al. 1993). Mean TL among 45 male and 51 female June

Suckers netted at Utah Lake in 1978-1979 = 50 cm and 55 cm, respectively (overall X ,--; 52 cm) (Radant and Sakaguchi 1981: 141). In both 1991 (n = 36) and 1992 (n = 56), mean TL of spawners = 53 cm for males and 57 cm for females (Gutermuth and Lentsch 1993: 8, Gutermuth et al. 1993: 8). Jordan (1878b),

Jordan and Gilbert (1881, 1882), Jordan (1891), and Jordan and Evermann (1896a) declared that with a TL of 30-33 cm and a mass normally < 0.5 kg, the Webug Sucker is the smallest year-round catostomid of

Utah Lake. Stubbs' (1966: tables 1-3) data corroborate that the BL of the Webug Sucker (X = 25 cm, n = 5) is appreciably < that of the Utah Sucker (X = 38 cm, n = 9) and June Sucker (X = 40 cm, n =

11), although Lowder (1951: tables 1-5) indicated that the BL of the Webug Sucker (X = 28 cm, n = 18) is > that of the Utah Sucker (X = 24 cm, n = 19) but < that of the June Sucker (X = 32 cm, n = 8).

The validity of Lowder's (1951) data on Utah Lake catostomids hinges on his ability to initially differentiate them correctly based principally on the use of mouth and head morphologies (see Stubbs 1966:

6). In identifying the Webug Sucker, Lowder (1951: 17-18) apparently relied heavily on the only published characterization of this species reproduced in his thesis. That description by Cope and Yarrow (1876: 678-

679) was based to a greater extent on specimens of the June Sucker than those of the Webug Sucker (see

Jordan 1878b: 219-220, Jordan et al. 1930: 108), and did not address mouthparts. Stubbs (1966: 7-8) noted that the configuration, papillosity, and plication of lips, which Lowder (1951) had largely employed to differentiate catostomids, are inconstant. Lowder (1951) did not refer to the characterizations of Utah Lake NATIVE PISCIFAUNA OF UTAH LAKE 49

suckers made by Jordan and Gilbert (1882: 128-129, 131-132) and Jordan (1891). Even though Lowder

(1951: 44) put "pp. 219" at the end of his citation of Jordan's (1878b) article, he actually quoted pages

149-150 and did not make reference to Jordan's (1878b: 219-220) brief portrayal of the Webug Sucker which

accompanied a restatement of the description of Chasmistes. Lowder (1951: 11) described only 1 ideal

representative specimen from each of 3 catostomid taxa. He did not statistically compare the 25

measurements and enumerations that he made on 45 collected specimens, and did not delineate any

morphometric, enumerative, or gross anatomical boundaries that he might have used in assigning a sucker

to 1 of the 3 species. Lowder (1951: 1, 9, 10, tables 1-5), who claimed to have inspected 6782 netted

suckers in 1950-1951 and to have performed complete measurements on =-- 100 of them (dimensions of 45

specimens are recorded), "recognized" near the end of his field survey of year-round Utah Lake catostomids

that there were 3 species. Lowder (1951: 1, 7, 9) repeatedly alluded to the presence of 3 suckers, and

devoted the body of his treatise to the presentation of morphologic data for each. Lowder (1951: 15-16)

affirmed that the Webug Sucker's short mandible, inferior mouth, and papillose lips clearly place it in

Catostomus. But Lowder (1951: 42) later attested that the catostomids exhibited a continuum of lip and head

traits that included all transitional levels of the features that had been described for these forms, concluding

"that there is either two genera and three species or one genus and one species of suckers in Utah Lake."

Then, Lowder (1951: 42) avowed that there must be 2 sucker species present because the June Sucker

spawns in April and the Utah and Webug Suckers spawn in June. The June Sucker derives its name from

the month during which its spawning activity has most often peaked (see Cope and Yarrow 1876: 678;

Jordan 1878b: 150-151; Jordan 1891; Stubbs 1966: 49; Radant and Sakaguchi 1981b: 144, 146-148, 150;

Radant and Shirley 1987; Radant et al. 1987: 1-2, 21, fig. 7; Modde and Muirhead 1990, 1994; Gutermuth

and Lentsch 1993; Gutermuth et al. 1993; Holden et al. 1994: 19, 27; USFWS 1995: 8). The Webug Sucker

also likely spawns in June (Cope and Yarrow 1876: 678-679, Jordan 1878b: 219-220). The Utah Sucker

breeds in March (Jordan 1891), late March and early April (Stubbs 1966: 49, Radant and Sakaguchi 1981:

144), April (Radant et al. 1987: 2), or April-May (Holden et al. 1994: 19). Reiterating the existence of a GREAT BASIN NATURALIST 50 continuous cline of sucker morphologic characters, Lowder (1951: 42-43) affirmed that the Webug Sucker is as different from its putative congener, the Utah Sucker, as the June Sucker is from the Utah Sucker.

There is a question whether Lowder (1951) accurately identified suckers caught in Utah Lake. In compliance with their description by Jordan (1878b: 149-150), Jordan and Gilbert (1881, 1882: 128-129,

131), and Jordan and Evermann (1896a: 179-183), his 3 representative specimens of the Utah Sucker, 0 Webug Sucker, and June Sucker had mandibles forming angles of 90° and 270°, 60° and 300°, and 45 and

315°, respectively, with the body's craniocaudal axis. However, Lowder (1951: 15-16) attested that in addition to its general appearance, the Webug Sucker's short mandible and inferior mouth, as well as its thick lips covered with spongy papillae, clearly place it in the genus Catostomus. Although the mouth of the

Webug Sucker is not big, its mandibles, similar to those of the June Sucker, are large and long (.-- 36% of

HL) (Jordan and Gilbert 1881, 1882: 129, Jordan 1891, Jordan et al. 1896a: 180-181). Since Jordan and

Evermann (1896b: 240, 1898b: 2794) and Jordan et al. (1930: 108) decided to place fecundus back in the genus Chasmistes, it may be assumed that the position of the Webug Sucker's mouth approximates that described for the genus (i.e., terminal) (Jordan and Gilbert 1882: 131, Jordan et al. 1896a: 182). More specifically, Jordan (1878b: 219) avowed that the conformations of the mouth and snout in the Webug and

June Suckers are "very similar". Furthermore, Cope and Yarrow's (1876) plate 32 (fig. 1), which supplied one basis for distinguishing the Webug Sucker (Jordan 1878b: 219-220), shows its mouth to be terminal.

Robert Rush Miller and Carl L. Hubbs had a notion that the Webug Sucker was a hybrid of the Utah and June Suckers (Sigler and Miller 1963: 164-165, Holden et al. 1974). Perhaps Miller and Smith (1981) attempted to confirm that suspicion. Their hypotheses have engendered uncertainty about the ancestry of the contemporaneous June Sucker and the legitimacy of the Webug Sucker as a distinct species, and have been accepted (usually uncritically) by most authors who have alluded to their monograph (Radant and Sakaguchi

1981a: 11, b: 135, 136, 139; Snyder and Muth 1988: 3; Modde and Muirhead 1990: 1; Gutermuth and

Lentsch 1993: 2-3; Holden et al. 1994: 19, 23; Utah Lake Fish Management Advisory Team 1994: 16, 31; but see U.S. Fish and Wildlife Service [USFWS] 1995: 1, 3). Miller and Smith (1981: 9, 11) postulated that NATIVE PISCIFAUNA OF UTAH LAKE 51

Catostomus fecundus was a hybrid of Catostomus ardens and Chasmistes liorus prior to a severe drought that continued from 1932-1935 and left Utah Lake almost dry during the winter of 1934-1935 (see Tanner

1936). They proposed that a self-perpetuating hybrid distinct from the apparently non-reproducing hybrid,

Catostomus fecundus, arose from the same parental Utah and June Suckers during the period of extreme low water in Utah Lake, but that this form belongs to a different genus than its pre-drought sister hybrid,

C. fecundus. The putative post-drought hybrid was named Chasmistes liorus mictus (Miller and Smith 1981:

15, 17). One of its parents, Chasmistes liorus liorus, is supposedly extinct; the other parent, Catostomus ardens, is extant. It is remarkable that partially on the basis of anatomical measurements of only 3 pre-drought and 9 post-drought specimens of the June Sucker, and the inspection of 6 exemplars of the

Webug Sucker, Miller and Smith (1981: 11, table 1) contended that the original form of the June Sucker no longer survives, that "it is unlikely that Catostomus fecundus ever existed as a distinct species in (Utah)

(L)ake", and that the existing June Sucker is a self-propagating intergeneral (not interspecific) hybrid. These authors did not refer to the investigations of Lowder (1951) and Stubbs (1966) involving at least the cursory inspection of thousands of Utah Lake suckers. Photographs presented in their report render some of their hypotheses suspect. The hippocrepiform mouth of a specimen of Miller and Smith's (1981) Chasmistes liorus mictus (Fig. 9C) resembles the hippocrepiform mouth of the single displayed specimen of C. 1. liorus

(Fig. 7A), rather than the lunate mouth of another supposed example of C. 1. mictus (Fig. 7B). Moreover, the mouth of the latter mictus appears smooth whereas the mouth of two other examples of putative mictus

(Figs. 9B and 9C) are papillose. In contrasting gill-raker counts, Miller and Smith (1981: table 3) examined larger samples of 14 supposed liorus and 32 mictus, as well as 18 C. ardens, and found little taxonomic overlap. Five photographs depicted the gill rakers of liorus to be rather amorphous, those of mictus to be compactly columnar, and those of fecundus to be sparsely columnar (Miller and Smith 1981: fig. 8).

Comparisons of 22 anatomical lengths in 3 specimens of pre-drought and 9 examples of post-drought June

Suckers revealed 3 dimensions that did not interspecifically overlap (Miller and Smith 1981: table 1). These measurements of predorsal length, HL, and ED were assessed in somewhat larger samples of 8 pre-drought GREAT BASIN NATURALIST 52 and 19 post-drought June Suckers, and showed little inter-sample overlap (Miller and Smith 1981: table 2).

With respect to the distance between the occiput and the origin of the dorsal fin, the greatest length recorded in 9 post-drought suckers was only 0.1% > the shortest length registered in 3 pre-drought suckers (31.5% of BL - represented by 2 pre-drought individuals) (Miller and Smith 1981: table 1). However, this length and predorsal length measure, to an appreciable extent, the same dimension. Miller and Smith (1981) performed a series of measurements and enumerations only on Utah Lake's pre- and post-drought June

Suckers. Except with respect to 1 trait, they did not compare putative Chasmistes 1. liorus and C. 1. mictus with Catostomus ardens to determine whether the dimensions or counts in mictus were intermediate. Also, they did not attempt to match specimens and recorded traits of putative C. 1. mictus with exemplars and recorded characteristics of the Webug Sucker to assess whether these forms might be synonymous. After measuring anatomic lengths, assessing the number and structure of gill rakers, counting scales, and analyzing head and mouth anatomies, Miller and Smith (1981), without performing statistical tests on their small samples, concluded that predorsal and caudal-peduncle lengths, mouth shape, lip papillosity, scale counts, and gill-raker shape and density, as well as head, mandibular, and ocular sizes in the putative ancestral and hybrid forms of the June Sucker are different. In totality, data on these forms from all sources (see also below) do not show a clear morphological difference; gill-raker counts are the most discriminating.

Lowder (1951: 41) attested that scale enumerations among the 3 year-round catostomids of Utah Lake completely overlap. The following data show, contrary to Miller and Smith's (1981: 15) determination, that most or all sets of recorded scale counts are too variable to be of taxonomic value in distinguishing Utah

Lake catostomids. The reported values of Su for pre-drought June Suckers were 60 (Cope and Yarrow 1876:

678-679; includes most of the specimens labeled C. fecundus; note that these authors' description was reiterated by Jordan [1878b: 219-220]), 61 (Jordan and Gilbert 1881), 63 (Jordan and Gilbert 1882: 132,

Jordan and Evermann 1896a: 183), 66 (Jordan 1891), but also 55-64 (Miller and Smith 1981: 12). In post-drought June Suckers, these quantities were 62-68 (.t = 64.4, n = 7) (Stubbs 1966: table 6), 60-70

(Miller and Smith 1981: 15), and 62-69 (.t = 65.2, n = 8) (Lowder 1951: table 1). Su in the Webug Sucker NATIVE PISCIFAUNA OF UTAH LAKE 53

= 60 (Cope and Yarrow 1876: 678-679 [only part of the specimens labeled this species belonged to it],

Jordan and Gilbert 1881, 1882: 129), 64 (Jordan and Evermann 1896a: 180-181), 64-70 (X = 67.4, n = 10)

(Stubbs 1966), but also 65-72 (X = 68.3, n = 18) (Lowder 1951: tables 4-5). In the Utah Sucker, Su = 65

(Jordan and Gilbert 1881, 1882), 61-69 (x = 66.1, n = 10) (Stubbs 1966), 63-71 (Jordan 1891), 70-72

(Jordan and Evermann 1896a: 179-180), but also 61-79 (X = 67.3, n = 18) (Lowder 1951: tables 2-3). S. in the pre-drought June Sucker = 7 (Jordan 1881), 9 (Jordan and Gilbert 1882: 132, Jordan and Evermann

1896a: 183), 9-11 (Miller and Smith 1981: 12), and 10 (Cope and Yarrow 1876: 678-679); S. in the post-drought June Sucker equaled 10-13 (X = 11.6, n = 8) (Lowder 1951: table 1) and 11-14 (X = 12.4, n = 12) (Stubbs 1966: table 5). In the Webug Sucker, S. = 8 (Jordan and Gilbert 1881, 1882; Jordan and

Evermann 1896a: 180-181), 10 (Cope and Yarrow 1876: 678-679), 10-12 (X= 11.1, n = 10) (Stubbs

1966), but also 11-14 = 12.3, n = 18) (Lowder 1951: tables 4-5). S. in the Utah Sucker = 9 (Jordan and Gilbert 1881, 1882: 128-129), 10-13 (X = 11.8, n = 10) (Stubbs 1966), 12 (Jordan and Evermann

1896a: 179-180), and 10-14 (X = 11.9, n = 18) (Lowder 1951: tables 2-3). SI, in the pre-drought June

Sucker = 8 (Jordan and Gilbert 1882: 132, Jordan and Evermann 1896a: 183), 9 (Jordan and Gilbert 1881),

8-10 (Miller and Smith 1981: 12), and 10 (Cope and Yarrow 1876: 678-679). In the post-drought June

Sucker, Sb = 8-11 (X = 9.2, n = 12) (Stubbs 1966: table 5) and 9-12 (X = 10.2, n = 8) (Lowder 1951: table 1). Sb in the Webug Sucker = 8 (Jordan and Gilbert 1881, 1882, Jordan and Evermann 1896a:

180-181), 8-9 (X = 8.8, n = 5) (Stubbs 1966), 10 (Cope and Yarrow 1876: 678-679), but also 10-14

(X = 11.3, n = 18) (Lowder 1951: tables 4-5). In the Utah Sucker, Sb = 8-10 (x = 9.1, n = 10) (Stubbs

1966), 9 (Jordan and Gilbert 1881, 1882: 128-129), 12 (Jordan and Evermann 1896a: 179-180), but also

9-14 (X = 10.9, n = 18) (Lowder 1951: tables 2-3). These data show that generally, scale counts were recorded to be lower by authors in the 1800s than by investigators in the 1900s. The value of Sil reported by Miller and Smith (1981: 12) for pre-drought June Suckers is relatively low. The magnitude and variability of Lowder's (1951: tables 1-5) Su, S., and Sb are relatively very high. GREAT BASIN NATURALIST 54

Lowder (1951: 41) related that D or A is not unique for any year-round Utah Lake catostomid, and

without evidence, maintained that the lengths of these fins are dependent on gender and reproductive

condition. Although Jordan and Gilbert (1881, 1882: 128-129, 132) and Jordan and Evermann (1896a:

179-183) concurred that A = 7 in the 3 catostomids, Lowder's (1951: tables 1-5) data show that A = 7 in

1 June Sucker and 8 in 7 others. In the pre-drought June Sucker, D = 11 (Jordan and Gilbert 1881, 1882:

132, Jordan and Evermann 1896a: 183), 12-13 (Cope and Yarrow 1876: 678-679 [most of the described

specimens belonged to this taxon]), and 13 (Jordan 1891). In the dorsal fin of the post-drought June Sucker,

Stubbs (1966: table 5) found 11-13 (mode of 12) rays and Lowder (1951: table 1) counted 12-13 (mode of

12) rays. The number of dorsal-fin rays in the Webug Sucker has been registered as 11 (Jordan and Gilbert

1881, 1882: 129), 11-12 (Jordan 1891, Jordan and Evermann 1896a: 180-181), 12-13 (Cope and Yarrow

1876: 678-679 [part of the exemplars labeled this species belonged to it], Lowder 1951: tables 4-5 [mode

of 12]), and 13-14 (mode of 13) (Stubbs 1966). In the Utah Sucker, D = 12-13 (Jordan and Gilbert 1881,

1882: 128-129; Jordan 1891, Jordan and Evermann 1896a: 179-180, Lowder 1951: tables 2-3 [mode of 12],

Stubbs 1966). Hence, fin-ray number is not useful in distinguishing the suckers of Utah Lake.

Noteworthily, not all of the traits of Lowder's (1951) Webug Sucker were intermediate between those

of the June Sucker and the Utah Sucker. For example, the basal length of the dorsal fin as a percentage of

BL was 16.3% (n = 18) in the Webug Sucker, 17.9% (n = 8) in the June Sucker, and 18.2% (n = 19) in

the Utah Sucker; the length of the anal fin's base as a percentage of BL was 7.6% (n = 18) in the Webug

Sucker, 8.2% (n = 19) in the Utah Sucker, and 9.0% (n = 8) in the June Sucker (Lowder 1951: tables 1-5).

Note, however, that Jordan and Gilbert (1881, 1882: 128-129, 132), Jordan and Evermann (1896a: 179-181,

183), and Stubbs (1966: 44-45) indicated that these fins are shortest in the June Sucker. Stubbs (1966: 49-50)

proclaimed that, of the > 1000 specimens belonging to 3 catostomid species that he examined, not one was

identified as an intermediate" although each taxon possessed some characters that were so. For example,

mouth and lip characters in the Webug Sucker, dorsal-rib morphology in the Utah Sucker, and the anteroposterior positioning of the articular process within the tripus (see below) of the June Sucker were NATIVE PISCIFAUNA OF UTAH LAKE 55 intermediate (Stubbs 1966). The treatment of catostomids given earlier in this section indicates that the

Webug Sucker is not intermediate in body mass, eye position, or the relative lengths of the body, head, and

P1. Other examples of non-intermediacy in this sucker are the shape and relative size of the dorsal spine complex, length and shape of the anterior ramus of the tripus, and length of the first vertebral centrum

(equaling 0.63% of BL in the Webug Sucker, 0.56% in the June Sucker, and 0.52% in the Utah Sucker

(Stubbs 1966). Since the June Sucker has a large, oblique mouth with smooth lips, and the Utah Sucker has a much smaller mouth with papillose lips (see above), it follows, assuming hybridism, that a moderately small and moderately oblique mouth (an intermediate condition) would have greater papillosity than a very large and oblique mouth. Notwithstanding, the lunate mouth shown in Miller and Smith's (1981) fig. 7B appears smoother than the larger hippocrepiform mouths in figs. 7C and 9C. Thus, the mouth of the putative example of mictus in fig. 7B is intermediate in shape but not in lip papillosity. Several more examples of morphologic non-intermediacy are revealed in Stubbs' (1966) study of principally internal traits of Utah Lake catostomids. Non-intermediate attributes for the Webug Sucker or Chasmistes liorus mictus are inconsonant with the position of Miller and Smith (1981) that these forms are hybrids of the June Sucker and Utah

Sucker. Miller and Smith (1981: 17) affirmed that "The hybrid origin hypothesis... would be rejected if the characteristics of mictus were inconsistent with hybrid-like intermediacy between Chasmistes liorus and

Catostomus ardens". Gel electrophoretic studies have demonstrated that a form that was presumed to be a hybrid of the Cut-ui (Chasmistes cujus) and Tahoe Sucker (Catostomus tahoensis) because of its intermediate mouth and head morphologies, is actually a variant of the Cui-ui (USFWS 1995: 3). This finding suggests that members of Chasmistes are pleomorphic or genetically polymorphic.

Stubbs (1966) attested that most external morphologic characters are ineffectual in differentiating the year-round catostomid residents of Utah Lake. Examples are scale density and the number of fin rays (see above). Stubbs (1966: 7) affirmed that in distinguishing Utah Lake suckers, it is important to consider a complex of characters, internal as well as external. Seeking anatomies that would more efficaciously distinguish these fishes, Stubbs (1966: 8) investigated the Weberian mechanism because its features are GREAT BASIN NATURALIST 56 unique in Catostotnidae, and had been used to define species as well as higher-level taxa within the family.

He discovered that there are many interspecific differences in the configuration and size of the components of the Weberian mechanism. Stubbs (1966: 42, 52) found that intTaspecific variability in the morphology of the Weberian mechanism is greatest in the relatively wide-ranging Utah Sucker and less in the Webug and

June Suckers which are endemic to the Utah Lake ecosystem. The conformations and sizes of the parts of the Weberian mechanism are most constant in the Webug Sucker.

The Weberian mechanism comprises the pars audit= (auditory part) which indirectly connects the swim bladder to the membranous labyrinth of the inner ear, and the pars sustentaculum (support part). The latter constitutes part of the anterior vertebral column that sustains and protects the pars audittun, and provides an attachment for muscles extending from the caudal cranium, pectoral girdle, cranial swim bladder, and vertebral elements that are caudad to the pars sustentaculum. The pars auditum is composed of four paired ossicles, the claustra, scaphia, tripi, and intercalaria (Stubbs 1966: 9). Claustra (Latin, claustrum, gateway, or a bolt or bar that secures a gate) are patelliform ossicles measuring 52.2 by 5.3.0 mm that lie at the anterior gateway to the Weberian apparatus (see Stubbs 1966: 21, plate 2). They articulate with the anterodorsal facet of the first vertebral centrum and the anteroventral margin of the second neural arch. The claustrum bears an articular condyle which Stubbs (1966: fig. 30) depicted graphically as being relatively longest in the Webug Sucker. Scaphia, measuring <3.6 by < 5.0 mm, extend craniocaudally from the claustra to the intervertebral space between the first and the second (comprising the fused second and third) centra. The scaphium, which in Latin means concavity (e.g., vessel or basin), derives its name from its cyathiform concha stapedis which is shallower in the Utah Sucker than the Webug and June Suckers. The scaphium bears (1) a dentate dorsal process that inserts in connective tissue at the ventrolateral juncture of the second and third neural arches, (2) a relatively large, obtuse, and peg-like ventral process that projects almost perpendicularly from the medial surface of the concha to insert into deep pits on the dorsolateral surface of the first vertebral centrum, and (3) a meager, blunt posterior process (Stubbs 1966: 22-23). The dorsal process is very acute and attenuated in the June Sucker, caniniform in the Utah Sucker, and gently NATIVE PISCIFAUNA OF UTAH LAKE 57

peaked in the Webug Sucker (Stubbs 1966: 22, 31, 37, plate 4). Contrary to Stubbs' (1966: plate 4)

graphical depiction of scaphia bearing a very short ventral process in the Webug Sucker and a more

prominent one in the June Sucker, the textual description of this ossicle (1966: 31) reveals that the ventral

process is larger and blunter in the Webug Sucker. Despite its much shorter appearance graphically, the

length of the ventral process in the Webug Sucker is 1.5 times the length of the dorsal process (Stubbs

1966: 22-23, fig. 31). In the Utah Sucker, the lengths of the two processes are approximately equal (Stubbs

1966: 37, fig. 22). From the posterior process which protrudes from the center of the lateral surface of each

concha stapedis, an interossicular ligament passes to the anterior ramus of the tripus and provides an abode

for the intercalaritun (Stubbs 1966: 23). Fenestrated tripi measuring <7.5 by 5 18.6 mm, continue caudally

and slightly ventrally from the dorsolateral surface of the first centrum to the mid-ventral part of the fifth

centrum (Stubbs 1966: 25, 32, 38). The tripus (three-foot: Latin tri-, three; Greek pous, foot) apparently

derives its name from three of its parts: (1) anterior ramus, (2) posterior ramus, and also the (3) articular

process that protrudes from the body (the fourth part of the tripus) both dorsally and ventrally, but is

continuous medially where it inserts into two dorsocaudally-directed grooves on the lateral surface of the

third vertebral centrum (Stubbs 1966: 25, plate 4). Of particular significance are the interspecific differences

in the configuration of the tripus from a lateral perspective. From this aspect, the rami are approximately

horizontal in the Webug Sucker (Stubbs 1966: 26, fig. 32); the anterior ramus is markedly decurved

(dorsally convex) in the June Sucker but recurved (dorsally concave) in the Utah Sucker; and the posterior

ramus is recurved in the June Sucker but decurved in the Utah Sucker (Stubbs 1966: 33, 39, figs. 25, 34).

The tripus possesses a dorsal fossa that is more craniad than caudad in position. In the Webug Sucker, the

base of this scalene dorsal fossa lies near the origin of the dorsal articular process (Stubbs 1966: 25, fig. 28).

The triangular dorsal fossa in the June Sucker is clearly distinguishable by its attenuated dorsal vertex which projects so far dorsally that it reaches well into the dorsal articular process (Stubbs 1966: 32, fig. 39). In

the Utah Sucker, the large, trapezoid dorsal fossa is differentiable because it protrudes cranially the greatest

distance. Ventrally, the tripi of the three year-round catostomids of Utah Lake can be discriminated easily. GREAT BASIN NATURALIST 58

In the Webug Sucker, there is a large, quadrangular ventral fossa that extends far into the posterior ramus, or two large fossae that are separated by a dividing brace that originates at the base of the ventral articular process (Stubbs 1966: 25, fig. 27). The bulk of the irregular ventral fossa in the June Sucker is situated in the body but is considerably prolonged into the posterior ramus (though not as much as in the Webug

Sucker) and extends well into the ventral articular process (this extension is not shown graphically) (Stubbs

1966: 32-33, fig. 38). Multiple small fossae exist craniad to the large ventral fossa of this species. In the

Utah Sucker, the shallow, triangular, ventral fossa is restricted to an area at, and slightly craniad to, the base of the articular process (Stubbs 1966: 38, fig. 19). The articular process exists only slightly craniad to the midpoint of the tripus in the Webug Sucker, much closer to the tip of the anterior ramus than to the tip of the posterior ramus in the June Sucker, and very close to the tip of the anterior ramus in the Utah Sucker

(Stubbs 1966: 25, 33, 38, plate 4). The anterior ramus of the tripus, which is connected to the scaphium via an interossicular ligament, is longest and most slender in the Webug Sucker (Stubbs 1966: 25-26, plate 4).

The base of the body of the intercalarium (intercalary ossicle) is embedded within or saddled over the mid-ventrocaudal portion of that ligament. Intercalaria measure <2.6 mm in length, and consist of a proximal body or manubrium (Latin, handle or haft) and a distal shaft that may bear a dorsal flange (Stubbs

1966: 24). In the Webug Sucker, the intercalaritun is a blocked ell whose caudal (outside) angle is almost perpendicular; this ossicle is broadly crescentic in the June Sucker and distinctively falcate in the Utah

Sucker (Stubbs 1966: plate 4). The shaft ascends from the manubrium to embed in the fibrous connective tissue covering the dorsolateral surface of the second vertebral centrum (Stubbs 1966: 24). Of systematic importance, according to Stubbs (1966: 38), the shaft in the Utah Sucker uniquely lacks a dorsal flange.

Distally, the shaft of the June Sucker is cochlear (spoon-shaped) (Stubbs 1966: 32).

With regard to the pars sustentaculum of the Weberian mechanism, the anatomy of vertebra 2 is interspecifically discriminating. Its neural arch, isolated from the corresponding centrum, is scutiform in the

Webug Sucker, ephippioid (saddle-shaped) in the June Sucker, and lachrymiform in the Utah Sucker (Stubbs

1966: 15, 28-29, 34-35, plates 1-2). In the Webug Sucker, dorsal ribs (also called transverse processes) that NATIVE PISCIFAUNA OF UTAH LAKE 59 are proximally wide project laterally from centrum 2 and also slightly dorsocaudally, becoming flattened and cultriform distally (Stubbs 1966: 15, plates 1-3). Two tiny apophyses on the proximal anterodorsal margin of the dorsal rib project medially to the mid-lateral surface of centrum 1 (Stubbs 1966: 15). The ventral surfaces of dorsal ribs in the Webug Sucker are broadly joined medially with the ventral (pleural) ribs of vertebra 2. Vertebra 3 lacks dorsal or ventral ribs (Stubbs 1966: 16). The pleural ribs of vertebrae 2 and

4 fuse in all 3 catostomids to form large wings (Stubbs 1966: 16, 29, 35, plates 1-3). The distance between the extremities of the dorsal ribs in the Webug Sucker is short, averaging =2/3 the width of its relatively squat 4th pleural ribs (Stubbs 1966: 16, plates 1-3). Dorsal ribs of the June Sucker are much more dorsocaudally angled than those of the Webug Sucker, and the distance between their extremities is much greater, averaging >90% of the distance between the termini of the 4th pleural ribs (Stubbs 1966: 16, plate

3). In this species, the dorsal rib is substantially fused ventrally with pleural rib 2, and the latter is joined with pleural rib 4 (Stubbs 1966: 29, plate 3). The configuration of the Utah Sucker's dorsal ribs (projecting laterad and only slightly caudad) and the distance between their extremities (g .--, 84% of the extreme width of the pleural ribs) is intermediate (Stubbs 1966: 35). Its dorsal ribs, in contrast to those of the other 2 suckers, are not as broadly fused with the 2nd pleural ribs, and the 2nd and 4th pleural ribs are not as extensively joined. In all 3 taxa, the pleural ribs of vertebra 2 extend laterally but also ventrocaudally and do not join medially on the ventral surface of centrum 2 (Stubbs 1966: 16, 31, 45). Contrariwise, pleural ribs of vertebra 4 fuse medially to form a transverse plate which lies ventrad to the mid-ventral surface of centrum 4 in the Webug and June Suckers but ventrad to the cranioventral portion of centrum 4 in the Utah

Sucker (Stubbs 1966: 20, 31, 36). From pyramidal cavities in the dorsolateral surfaces of centrum 4, pleural ribs extend ventrolaterally in the Utah and Webug Suckers but ventrocaudally in the June Sucker (Stubbs

1966: 19-20, 30-31, 36, plates 1-3). Prominent ridges existed on the pleural ribs except in their craniad portion. The mean distance between the distal extremities of the 4th pleural ribs = 11.35% of mean BL in the Webug Sucker, 11.64% in the Utah Sucker, and 11.78% in the June Sucker (Stubbs 1966: derived from tables 1-4). The massive neural spine complex of vertebra 3 projects caudad and is cultriform dorsally GREAT BASIN NATURALIST 60

(Stubbs 1966: 17). Overall, it is scapuloid and caudally acute in the Webug Sucker, sagittate in the June

Sucker, and broadly sagittate in the Utah Sucker (Stubbs 1966: plates 1-2). In the Webug Sucker this complex is relatively short, extending caudad to a point midway between neural spines 6 and 7; it continues caudally to the apex of neural spine 7 in the June and Utah Suckers (Stubbs 1966: 17, 29, 35).

June Sucker Chasmistes 'fonts (smooth-bordered yawner, in reference to its large, oblique gape with lips devoid of papillae: Latin ho, to smooth [Greek leios, smooth]; Greek oros, border or margin). Cope and

Yarrow's (1876) description of Catostomus fecundus was based on a combination of specimens belonging to both Catostomus fecundus and Chasmistes liorus (see Jordan 1878b). As Jordan (1878b: 219-220) explained in an addendum, the form described under the rubric of Chasmistes fecundus on pages 150-151 of the same treatise, is actually C. liorus. There are 37-53 (.7z 44) gill rakers on the first branchial arch of the June Sucker (Miller and Smith 1981: see table 3). In males, the anal fin and the lower lobe of C are tuberculate (Jordan and Gilbert 1881). The parietal fontanelle is very large (Jordan 1878b, Jordan and Gilbert

1882, Jordan and Evermann 1896a). Other characteristics of the June Sucker are compared with those of other year-round catostomids of Utah Lake in the section on the Webug Sucker (see above).

ORDER SALMONIFORMES

Family Salmonidae

Mountain Whitefish Prosopium williamsoni (Williamson's masked one: Greek prosopion, mask [referring to the species' large preorbital bones]; williamsoni refers to Lieutenant R. S. Williamson who commanded the U.S. railroad survey of California and Oregon [Girard 1857b]). This salmonid was referenced by the binomen Coregonus williamsoni (Girard 1857b, 1858b: 326-327) (C. williamsonii of Cope [1872a] and

C. villiamsonii of Cope [1872b, 1876, 1879] and Cope and Yarrow [1876: 638, 682-683]) until Jordan

(1928) placed it in the genus Prosopium. Prosopium couesii (Milner 1874, Jordan 1878c: 276, 362, Jordan and Gilbert 1882: 297; see also Jordan and Evermann 1896a: 463 and Bailey et al. 1948: 8) is a putative synonym. This wide-ranging species (Sigler and Sigler 1987: 106-108, Page and Burr 1991: 46-47) was reported to be "very abundant" in Utah Lake {Jordan and Gilbert 1881). Jordan (1891) signified that it was NATIVE PISCIFAUNA OF UTAH LAKE 61 seldom encountered in the lake but was common in Provo River a short distance upstream from Provo. The

Mountain Whitefish was taken at Utah Lake near Provo River's mouth in March 1927 (Tanner 1936).

Prosopium williamsoni has a BD/TL that = 20.7-23.5%, a small, subconic, and short (HL/TL =

18.2-22.2%) head with small eyes (ED/HL = 16.7-21.4%) and mouth, as well as a compressed, protrusive, and subacute or obtuse snout lying below eye level (Girard 1857b, 1858b: 326-327, Cope and Yarrow 1876:

682-683, Jordan and Gilbert 1882: 297-298, Jordan 1891). Along williamsoni's straight lateral line, Su =

74-88; S. = 8-10 and Sb = 7-10; D = 11-14 and A = 10-12 (Girard 1857b, 1858b, Cope and Yarrow

1876, Jordan and Gilbert 1882). In P. couesii, BD/TL and HL/TL both = 22.5-24.2%, Sil = 88, S. = 8,

Sb = 5-8, D = 12, and A = 10, approximating corresponding measurements and counts in P. williamsoni; however, in couesii, the head tapered rostrad gradually to the point of union of the supraorbital bones, whence it narrowed abruptly, terminating in a extremely protuberant snout that was porcine in the breeding male (Milner 1874, Jordan and Gilbert 1882, Jordan and Evermann 1896a: 463). Hence, couesii may be a distinct form. Erroneously, Jordan (1878c) declared that in couesii, D = 111-12, A = 1V-10, and Su = 38.

In the Mountain Whitefish, heights of the slightly concave dorsal and anal fins are > basal lengths

(Girard 1858b: 326-327). Scales are tuberculate on the sides of breeding males; the large adipose fin extends farther caudad than the anal fin; P1 are short, and P2, which originate ventrad to the caudal 'A of the dorsal-fin base, proceed caudad to a point that is far short of the vent (Girard 1858b, Jordan and Gilbert

1882: 297-298, Jordan and Evermann 1896a: 463). Broad and short (length ---z 'A of HL) maxillae extend no farther caudad than the rostral margin of the eyes; mandibular length = Vs of HL (Girard 1857b, 1858b,

Cope and Yarrow 1876: 682-683, Jordan and Gilbert 1882, Jordan and Evermann 1896a). williamsoni has a slaty dorstun, and silvery white sides and belly (Girard 1857b, 1858b, Cope and Yarrow 1876). According to Jordan and Evermann (1896a), caudal and adipose fins are steely blue, and all fins are tipped with black.

TL = 432-457 mm in examples taken from Provo River in 1872 (Cope and Yarrow 1876), and 190-425 mm

(mass = 250-600 g) in 74 samples obtained from Provo and Weber Rivers in 1970 (Hansen 1970).

Bonneville Cutthroat Trout Oncorhynchus clarki utah (Clark's bulky-snouted Utahn: Greek Onkos , bulky; rhunkhos , snout; clarki refers to Captain William Clarke (1770-1838) of the famed Lewis and Clarke GREAT BASIN NATURALIST 62 expedition; in the original published account of the exploration [Allen 1814], Clarke's name was spelled

"Clark"). Cope (1872b) indicated that Girard's (1857a: 26, d, 1858b: 320-321) Salar (=Salmo) virginalis in which BD / the distance from the tip of the snout to the notch of C = 17.4% and ED/HL = 22.2%, was distinct from Cope's (1872a) Salmo (Salar) virginalis in which BD / the distance from the tip of the snout to the notch of C = 22.2% and ED/HL = 25%, and assigned the name Salmo pleuriticus to the latter which existed in Henry's Fork of Green River about the border of Wyoming and Utah. BD / the distance from the snout's tip to the notch in C = 19.0% in Saltily carinatus and 21.0% in S. spilurus (Cope 1872b). Cope

(1876) declared that the trout from Utah Lake was Salmo virginalis because it was slimmer (BD/TL =

16.7-17.4%) than S. pleuriticus whose BD/TL = 21.5%. Yarrow (1876) also referenced the trout of Utah

Lake as S. virginalis. Cope and Yarrow (1876: 685-693) did not regard S. pleuriticus as distinct from Salmo virginalis. The cutthroat trout from Utah was called Salar spilurus by Jordan (1878a: 431), Salnw stomias spilurus and S. clarkii by Jordan (1878c: 358-359), and Salmo purpuratus by Jordan and Gilbert (1881).

Jordan and Gilbert (1882: 314-315) affirmed that S. spilurus comprised fish from the "Basin of Utah" as well as the "upper Rio Grande", and regarded S. pleuriticus of Cope (1872b) as a subspecies. Also, they lumped Salmo carinatus, S. ciarki (Richardson 1836), S. utah (Suckley 1874: 135-138 [based on a memoir delivered to the Smithsonian Institution in 1861]), and Salar virginalis within Salmo purpuratus . Then Jordan

(1886a) included spilurus and virginalis as subspecies of S. purpuratus, noting that virginalis was endemic to "Utah Basin", but that spilurus was restricted to "Rio Grande and Colorado Basin." He admitted that he could not differentiate Utah Lake examples of S. purpuratus from exemplars in Columbia River. In contradistinction, Jordan (1887a) maintained that S. purpuratus and S. spilurus were both full species from

Salt Lake Basin. Jordan (1887b: 832) made stomias and spilurus subspecies of S. purpuratus, but omitted virginalis as a nominal form. Then Jordan (1891) and Jordan and Evermann (1896a: 495, b: 291) declared that Salmo rnykiss virginalis comprised both Salar virginalis (Girard 1857a,d, 1858b) and Salmo utah

(Suckley 1874), but were under the erroneous impression that S. virginalis had been taken from Utah Lake or its tributaries. Jordan and Evermann (1902: 182) regarded Salmo virginalis as a full species. Finally,

Jordan (1920) realized that Salar virginalis was not obtained in Utah and made Suckley's (1874) Salmo utah NATIVE PISCIFAUNA OF UTAH LAKE 63 the name of cutthroat trout inhabiting Utah's Wasatch Front. Jordan (1920) was mistaken in noting that Utah

(=Ute) Creek (of Costilla County in southern Colorado) is located near Wagonwheel Gap. Rather, it originates on the northeast margin of present-day Mount Lindsey in the Sangre de Cristo Mountains and proceeds south-southwestward in a valley, passing Little Bear Peak to its west, then Buck Mountain to its east, and joining Trinchera Creek outside Fort Garland (U.S. Department of the Interior, Geological Survey

[1973], State of Colorado [map]; compare C. Roeser [1879], State of Colorado [map], U.S. Department of the Interior, General Land Office). In the 1850s, Fort Massachusetts, situated on Utah Creek's bank 8 km upstream (north-northeast) from Fort Garland (New Sectional Map of Colorado [1889], Rand, McNally &

Co., Publishers, Chicago) was operational; Captain J. W. Gunnison's exploratory party was camped a short distance below that fort when Salar virginalis was taken (Beckwith 1854: 38-39, 41). Girard (1857d) asserted that Salar virginalis was acquired in both Utah Creek and Sangre de Cristo Pass whereas Girard (1857a,

1858b: 320-321) recorded only the collection site of Utah Creek for this form. "Considerable" labor by a large workforce was expended during 4 days to construct a crude road > 19 km in length within Sangre de

Cristo Pass (Beckwith 1854: 38-39). Gunnison's party diverged from Sangre de Cristo Creek --.. 13 km below the summit of that pass and camped on Utah Creek > 17 km below the summit for several days. If, as seems likely, arduous road-building did not allow time for catching fish, F. Creutzfeldt's trout collections were probably confined to the 3 examples taken in Utah Creek (Girard 1858b). Notwithstanding, he could have acquired exemplars in Sangre de Cristo Pass either while wagons of his party were passing through it or while he was camped at Utah Creek. Jordan (1928) regarded the Bonneville Cutthroat Trout as the subspecies S. clarki utah, but Jordan et al. (1930: 56-57) and Tanner (1936) gave S. utah full-species status.

Schrenkeisen (1938: 44) called this form S. clarkii utah. La Rivers and Trelease (1952) considered the spelling to be S. clarki utah. Smith and Stearley (1989) showed that cutthroat trout are in the genus

Oncorhynchus. Robins et al. (1991) sanctioned the binomen 0. clarki, and Stearley and Smith (1993) confirmed its appropriateness. The Utah subspecies is 0. clarki utah (Utter and Allendorf 1994).

In the Bonneville Cutthroat Trout, BD/TL = 16.7-17.4%, HL/TL ,----, 23%, D = A = 12, predorsal length is slightly <50% of BL, P2 originate ventrad to ray 6 of the dorsal fin, Su = 145-180, S. = 36-42, GREAT BASIN NATURALIST 64 and SI, = 40-41 (Suckley 1874: 135-138, Cope and Yarrow 1876: 685-693, Jordan 1891, Holden et al.

1974). Maxillae proceed caudad to a point 74=5 mm beyond the posterior margin of the eyes, the labial commissure exists ventrad to the caudal border of the pupil, and C is slightly forked. Suckley (1874) stated that in Utah Lake examples, the fulvous dorsum was speckled black above the straight lateral line. He added that sparse dorsal stippling existed craniad to the origin of the yellowish dorsal fin, larger stellate markings occurred in the dorsum caudad to the origin of the dorsal fin and craniad to the anus, and yet larger, dense splotches were exhibited caudad to the anus both above and below the lateral line. Jordan (1891) noted that large Utah Lake trout were deep green with silvery sides, faint dark speckles, yellowish dorsal fins, and red ventral fins. According to Suckley (1874), ventral fins were rosy with red rays. Small oval or round black spots appeared on the dorsal, adipose, and caudal fins, on the opercula and atop the head. The silvery white venter was separated from the dorstun by a faint, irregular golden band. Suckley (1874) related that ventral vermilion streaks 3 mm wide extended caudad from the white chin to a point ventrad to the middle of the opercula. Holden et al. (1876) characterized these markings as lemon-yellow slashes in live individuals. The iris was golden bronze with circular black spots the size of a pinhead (Suckley 1874). Yarrow (1876) noted that during the spawning period, the coloration of the male, but particularly of the female, became more brilliant due at least partially to the increased engorgement of superficial blood vessels. The luster of the female increased more than that of the male during the breeding season, especially in the lower cheek. At this time, the female enlarged, the smaller male was either more (p. 364) or less (p. 363) radiant than the female, and in both sexes, the eyes became brighter and the typical sluggishness ended (Yarrow 1874).

In 1851, 1 trout from Timpanagos (=Provo) River had a TL of 660mm and a mass of =2.5 kg, and in 1860, the largest 2 of 3 individuals obtained there weighed 0.8 kg and 1.0 kg (Suckley 1874: 135-138).

Lieutenant R. S. Williams caught an individual in that river =11 km upstream from Utah Lake which weighed 3.2 kg (Suckley 1874). Maximal TL in this subspecies = 762 mm according to Suckley (1874) but

915 mm according to Yarrow (1874) and Cope and Yarrow (1876: 685-693); maximal mass = 7.0 kg

(Yarrow 1874, Cope and Yarrow 1876). In 1872, mean TL and mass 2--, 356 mm and 0.7 kg, respectively

(Yarrow 1874). TL = 368 and 386 mm in 2 specimens that were collected in 1872 (Cope and Yarrow 1876) NATIVE PISCIFAUNA OF UTAH LAKE 65 and 380 mm in an individual that was netted in 1959 (Arnold 1959). In 1889, one seine retrieval in a channel of Utah Lake's Goshen Bay yielded 50 trout whose mass ranged from 0.9-1.6 kg (Jordan 1891).

ORDER SCORPAENIFORMES

Family Cottidae

Prickly Sculpin Cottus semiscaber (semi-rough sculpin: Greek kottos, sculpin; Latin semiscaber, semi-rough). Bailey and Bond's (1963) Cottus echinatus from Utah Lake is a valid species only if it is distinct from Cottopsis semiscaber which was collected in the Pocatello, Idaho area (Fort Hall) (Cope 1872b; see Jordan 1878a: 441). Although it may be separate, Bailey and Bond (1963) did not compare the 2 forms or even report on the 3 specimens from Fort Hall, but simply proclaimed that Cope's (1878b) semiscaber is "properly a specific synonym of bairdi." The first statement in the species description of C. echinatus by Bailey and Bond (1963) reads, "Jordan and Gilbert (1881) and Jordan (1891: 35) noted the prickly skin of this species". In actuality, Jordan and Gilbert (1881) declared, "Our (2) specimens are villous above and below" the lateral line. Bailey and Bond (1963) did not mention the "villous" skin of Jordan and Gilbert's

Utah Lake specimens; a determination about their villosity versus spinosity must be made. Those 2 exemplars differed from C. semiscaber whose skin was "prickly above the lateral line, (but) smooth below it posteriorly" (Cope 1872b), Uranidea wheeleri from Utah's Beaver River whose skin was "everywhere smooth" (Cope 1876), but also from Jordan's (1891) prickly-skinned examples from Utah Lake. Nonetheless,

Jordan and Gilbert (1882: 695) and Jordan (1887b: 898) synonymized Utah Lake and Fort Hall sculpins, calling them Uranidea semiscabra. Jordan (1891) also considered their Cottus semiscaber to comprise sculpins from both sites, referring the reader to Jordan and Gilbert (1882) for the species description, a characterization based on both forms. Bailey and Bond (1963) wrongly signified that Jordan's (1891) notation about prickly skin applied only to the sculpin of Utah Lake. Jordan and Evermann (1898a: 1949-1950) subsumed the smooth-skinned Uranidea wheeleri and Gill's (1862a, 1876) Potamocottus punctulatus, which

Jordan (1891) regarded as a synonym of U. wheeleri, within C. semiscaber. Jordan and Evermann (1896b:

440), Jordan et al. (1930: 384), and Tanner (1936) gave P. punctulatus full-species status but considered the

Fort Hall and Utah Lake forms as well as U. wheeleri to belong to C. semiscaber. Their C. semiscaber was GREAT BASIN NATURALIST 66 regarded as a subspecies of C. bairdi by Sigler and Miller (1963), and apparently even earlier by Bailey et al. (1960). Robins etal. (1991: 41) accepted Bailey and Bond's (1963) name of C. echinatus for supposedly endemic Utah Lake sculpins. Sculpin specimens that have been collected in Utah Lake need to be carefully compared with Cope's (1878b), and if possible, future exemplars from waterways in Bannock County, Idaho.

Unfortunately, Utah Lake sculpins are believed to have been extirpated (Robins et al. 1991).

Bailey and Bond (1963) affirmed that Utah Lake harbored an endemic prickly-skinned sculpin species where 2 exemplars were obtained in 1880 (Jordan and Gilbert 1881) and 2 examples were supplied to the

U.S. National Museum by Peter Madsen in 1882. The holotype and 2 paratypes of Bailey and Bond's (1963)

C. echinatus were taken in Utah Lake near Provo River's mouth in April 1928 by Vasco M. Tanner, and furnished to the University of Michigan Museum of Zoology. Although the uniqueness of this supposedly new species rests on the relative spinosity of its skin, Jordan and Evermann (1898a: 1949-1950) attested that there is much variability in skin roughness among sculpins from Utah and southern Idaho. Bailey and Bond

(1963) noted, "So far as known Cottus echinatus is restricted to Utah Lake, Utah", but added that although

Jordan (1891) reported it in Provo River, the collection site of those examples of the taxon in the U.S.

National Museum that were collected by Jordan was Utah Lake. That those observed sculpin specimens were from Utah Lake does not nullify Jordan's (1891) report that semiscaber was "not rare" in Provo River.

With respect to the measurement and counts of anatomical parts, there was no appreciable difference between semiscaber from Fort Hall (Cope 1872b) and echinatus from Utah Lake (Bailey and Bond 1963) except that ED / interorbital width supposedly = 1% in the former but 2.7 in the latter. D = VI1,18 in the former and VII-VI11,16-18 in the latter, A = 13 in the former and 13-14 in the latter, HL/BL = 33% in the former and 31-35% in the latter, and ED/HL = 22.2% in the former and 23.6% in the latter. Jordan and Evermann (1898a: 1949-1950) signified that sculpin examples from "southern Idaho (Fort Hall) and

Provo River, Utah" had the roughest skin due to a bounty of slender papillae. In sculpins from Utah Lake and Fort Hall, the cuneate head was depressed dorsomedially, the mouth was large, and the maxillae extended caudad to a point ventrad to the posterior part of the pupil (Jordan and Gilbert 1882: 695, Bailey and Bond 1963). BL = 61-92 mm, TL < 110 mm, BD/BL = 19.0-23.5%, depth of the caudal peduncle NATIVE PISCIFAUNA OF UTAH LAKE 67

/ BL = 6.6-7.4%, P1 approached the anal fin caudally, and P2 extended caudally only ih the distance to the anal fin. In sculpins from Utah Lake, SL/BL = 9.4%, predorsal length = 36.7% of BL, prepelvic length

= 29.7% of BL, and caudal-peduncle length = 17.2% of BL (Bailey and Bond 1963). Anal-fin basal length

(29.1% of BL) was much > depth (13.4% of BL); likewise, the spiny and soft dorsal fins were much longer

(lengths = 19.5% and 40.6% of BL, respectively) than high (heights = 9.4% and 17.0% of BL, respectively). The gape was oblique, palatine teeth were well developed, and jaws were straight and equal; in ventral aspect, the upper jaw was only narrowly visible. The low, spinous fin of Utah Lake sculpins barely joined or was separate from the higher soft dorsal fin that continued caudad almost to the base of the long C (Bailey and Bond 1963). Jordan and Gilbert (1882) noted that the entire body except the ventral caudal peduncle was tuberculate, but that only tubercles of the anterior and dorsal parts of the body bore a spine. According to Bailey and Bond (1963), skin prickles in Utah Lake sculpins existed on the body as far caudad as the posterior end of the soft dorsal fin. Cephalic spines of these fish had a median pore about the size of the naris. Jordan and Gilbert (1882) affirmed that Utah Lake and Fort Hall sculpins had opercular and preopercular spines. Bailey and Bond (1963) did not mention "opercular" spines, but declared that there were either 4 complete and sharp preopercular spines, or 3 preopercular spines which were complete and

1 that was knobby; the dorsal preopercular spine was either straight or recurved, and the 2nd spine was directed ventrocaudally rather than being hooked ventrocranially as it is in other American members of the genus, with 1 exception. In the Utah Lake sculpin, the straight but incomplete lateral line, lying somewhat dorsad to the axial septum, bore 27 pores and terminated caudally at a point ventrad to soft dorsal rays 16-18

(Bailey and Bond 1963). The dorsum was tawny, the venter was pallid, the sides were faintly mottled, and

2-3 dark markings existed at the base of C. In males, the spinous dorsal fin was darkly pigmented both cranially and caudally. Bold cross bands appeared on the soft dorsal and caudal fins, and faint markings occurred on the pelvic and anal fins of males (Bailey and Bond 1963). GREAT BASIN NATURALIST 68

LITERATURE CITED

ALLEN, P. (editor). 1814. History of the expedition under the command of Captains Lewis and Clark, to

the sources of the Missouri, thence across the Rocky Mountains and down the River Columbia to the

Pacific Ocean. Bradford and Inkskeep and Abraham H. Inskeep, New York. 1: 1-470, 2: 1-522.

ARNOLD, B. 1959. Monthly report for April, 1959. Unpublished document, Utah Department of Natural

Resources, Division of Wildlife Resources. 17 pp. + 1 figure.

ARNOLD, B. B. 1960. Life history notes on the Walleye, Stizostedion vitreum vitreum (Mitchill) in a turbid

water, Utah Lake, Utah. Unpublished master's thesis, Utah State University, Logan. 107 pp.

AYRES, W. 0. 1854. (Untitled). Proceedings of the California Academy of Natural Sciences 1: 20-21.

BAILEY, R. M., AND C. E. BOND. 1963. Four new species of freshwater sculpins, genus Cottus, from

western North America. Occasional papers of the Museum of Zoology, University of Michigan

No. 634: 1-27.

BAILEY, R. M., E. A. LACHNER, C. C. LINDSEY, C. R. ROBINS, P. M. ROEDEL, W. B. SCOTT,

AND L. P. WOODS. 1960. A list of common and scientific names of fishes from the United States and

Canada. Second ed. American Fisheries Society Special Publication No. 2: 1-102.

BAIRD, S. F., AND C. GIRARD. 1854a. Descriptions of some new fishes from the river Zuni. Proceedings

of the Academy of Natural Sciences of Philadelphia (1852, 1853) 6: 368-369.

- --. 1854b. Descriptions of new species of fishes collected by Mr. John H. Clark, on the U.S. and

Mexican boundary survey, under Lt. Col. Jas. D. Graham. Proceedings of the Academy of Natural

Sciences of Philadelphia (1852, 1853) 6: 387-390.

- --. 1854c. Description of new species of fishes, collected by Captains R. B. Marcy, and Geo. B.

M'Clellan in Arkansas. Proceedings of the Academy of Natural Sciences of Philadelphia (1852, 1853) 6: 390-392.

- --. 1856. Descriptions of new species of fishes collected in Texas, New Mexico and Sonora, by Mr. John

H. Clark, on the U.S. and Mexican boundary survey, and in Texas by Capt. Steward Van Vliet, U.S.A.

Proceedings of the Academy of Natural Sciences of Philadelphia (1854, 1855) 7: 24-29. NATIVE PISCIFAUNA OF UTAH LAKE 69

BARBOUR, C. D., AND R. R. MILLER. 1978. A revision of the Mexican cyprinid fish genus Algansea.

Miscellaneous Publications, Museum of Zoology, University of Michigan No. 155: 1-72.

BECKVVITH, E. G. 1854. Report of exploration of a route for the Pacific railroad, near the 38th and 39th

parallels of latitude, from the mouth of the Kansas to Sevier River, in the Great Basin. Unpublished

report. 103 pp + 1 map.

CARBINE, W. F. 1936. The life history of the chub, Tigoma atraria Girard, of the Great Basin area of Utah.

Unpublished master's thesis, University of Utah, Salt Lake City. 107 pp. + map and 5 plates.

CARTER, D. R. 1969. A history of commercial fishing on Utah Lake. Unpublished master's thesis, Brigham

Young University, Provo, UT. 142 pp.

CHU, Y. T. 1935. Comparative studies on the scales and on the pharyngeals and their teeth in Chinese

cyprinids, with particular reference to taxonomy and evolution. Biological Bulletin of St. John's

University No. 2: 1-225 + 30 plates.

CHUTE, W. H., R. M. BAILEY, W. A. CLEMENS, J. R. DYMOND, S. F. HILDEBRAND, G. S.

MYERS, AND L. P. SCHULTZ. 1948. A list of common and scientific names of the better known

fishes of the United States and Canada. American Fisheries Society Special Publication No. 1: 1-45.

COBURN, M. M., AND T. M. CAVENDER. 1992. Interrelationships of North American cyprinid fishes.

Pages 328-373 in R. L. Mayden, editor, Systematics, historical ecology, and North American freshwater

fishes. Stanford University Press, Stanford, CA. 969 pp.

COPE, E. D. 1864. Partial catalogue of the cold-blooded Vertebrata of Michigan. Proceedings of the Academy

of Natural Sciences of Philadelphia (1864) 16: 276-285.

---. 1872a. Recent reptiles and fishes: report on the reptiles and fishes obtained by the naturalists of the

expedition. Pages 432-442 in F. V. Hayden, editor, Preliminary report of the United States Geological

Survey of Wyoming, and portions of contiguous territories, (being a second annual report of progress)

(1870). Government Printing Office, Washington, DC. 511 pp.

1872b. Report on the recent reptiles and fishes of the survey, collected by Campbell Carrington and

C. M. Dawes. Pages 467-476 in F. V. Hayden, editor, Preliminary report of the United States GREAT BASIN NATURALIST 70

Geological Survey of Montana and portions of adjacent territories; being a fifth annual report of progress

(1871). Government Printing Office, Washington, DC. 538 pp.

- --. 1876. On the Plagopterinae and the ichthyology of Utah. Proceedings of the American Philosophical

Society (1874-1875) 14: 129-139.

---. 1879. A contribution to the zoology of Montana. American Naturalist 13: 432-441.

- --. 1884. On the fishes of the Recent and Pliocene lakes of the western part of the Great Basin, and of

the Idaho Pliocene lake. Proceedings of the Academy of Natural Sciences of Philadelphia (1883) 35:

134-167.

COPE, E. D., AND H. C. YARROW. 1876. Report upon the collections of fishes made in portions of

Nevada, Utah, California, Colorado, New Mexico, and Arizona, during the years 1871, 1872, 1873, and

1874. Report Upon Geographical and Geological Explorations and Surveys West of the One Hundredth

Meridian, in Charge of First Lieut. Geo. M. Wheeler, Corps of Engineers, U.S. Army, under the

direction of Brig. Gen. A. A. Humphreys, Chief of Engineers, U.S. Army 5: 635-703 + plates 26-32.

CRAWFORD, M. 1979. Reproductive modes of the least chub (lotichthys phlegethontis - Cope). Unpublished

master's thesis, Utah State University, Logan. 79 pp.

CUVIER, LE BON, AND A. VALENCIENNES. 1842. Histoire naturelle des poissons. Vol 16. P. Bertrand,

Paris, France. 472 pp.

EVERMANN, B. W. 1893. A reconnaissance of the streams and lakes of western Montana and northwestern

Wyoming. Bulletin of the United States Fish Commission (1891) 11: 3-60 + plates 2-26.

GARMAN, S. 1880-1881. New and little-known reptiles and fishes in the museum collections. Museum of

Comparative Zoology at Harvard College, in Cambridge 8: 85-93.

---. 1881. North American fresh water fishes. Science Observer 3: 57-60.

GILL, T. 1862a. Observations on the genus Cottus, and descriptions of two new species (abridged from the

forthcoming report of Capt. J. H. Simpson). Proceedings of the Boston Society of Natural History

(1861-1862) 8: 40-42. NATIVE PISCIFAUNA OF UTAH LAKE 71

- --. 1862b. Description of a new species of the genus Tigoma of Girard (abridged from the forthcoming

report of Capt. J. H. Simpson). Proceedings of the Boston Society of Natural History (1861-1862) 8: 42. ---. 1876. Report on ichthyology. Pages 383-431 and plates 1-9 in J. H. Simpson, editor, Engineer

Department, U.S. Army: report of explorations across the Great Basin of the Territory of Utah for a

direct wagon-route from Camp Floyd to Genoa, in Carson Valley, in 1859. Government Printing Office,

Washington DC. 518 pp. GIRARD, C. 1857a. Report on fishes collected on the survey. Pages 21-27 and plates 7, 23, 49, 54, 56, 61,

and 73-75 in E. G. Beckwith, editor, Zoological report, which in turn was in E. G. Beckwith, editor (dated 1854), Report of Lieut. E. G. Beckwith, Third Artillery, upon explorations for a railroad route,

near the 38th and 39th parallels of north latitude, by Captain J. W. Gunnison, Corps of Topographical

Engineers, and near the forty first parallel of north latitude, by Lieut. E. G. Beckwith, Third Artillery.

Explorations and Surveys for a Railroad Route from the Mississippi River to the Pacific Ocean. War

Department, Washington, DC. Vol. 2, Part 4.

- --. 1857b. Contributions to the ichthyology of the western coast of the United States, from specimens in

the museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of

Philadelphia (1856) 8: 131-137.

- --. 1857c. Researches upon the cyprinoid fishes inhabiting the fresh waters of the United States of

America, west of the Mississippi Valley, from specimens in the museum of the Smithsonian Institution.

Proceedings of the Academy of Natural Sciences of Philadelphia (1856) 8: 165-213.

- --. 1857d. Notice upon the species of the genus SaImo, of authors, observed chiefly in Oregon and

California. Proceedings of the Academy of Natural Sciences of Philadelphia (1856) 8: 217-220. - --. 1858a. Notice upon new genera and new species of marine and fresh-water fishes from western North America. Proceedings of the Academy of Natural Sciences of Philadelphia (1857) 9: 200-202. ---. 1858b. Fishes. Explorations and surveys for a railroad route from the Mississippi River to the Pacific Ocean - War Department 10(4): 1-400 + 76 plates. GREAT BASIN NATURALIST 72

GUTERMUTH, F. B., AND L. D. LENTSCH. 1993. Reproductive biology studies of the June Sucker in the

Provo River, Utah: Draft 1991 progress report. Utah Department of Natural Resources, Division of

Wildlife Resources, Salt Lake City. 26 pp.

GUTERMUTH, F. B., L. D. LENTSCH, AND MARK C. STANGER. 1993. June Sucker reproductive

biology in the Provo River, Utah: Draft 1992 progress report. Utah Department of Natural Resources,

Division of Wildlife Resources, Salt Lake City. 19 pp. + Appendix 1 (2 pp.).

HANSEN, F. R. 1970. A study of the electrophoretic patterns of blood serum proteins from two populations

of Mountain Whitefish (Prosopium williamsoni). Unpublished master's thesis, Brigham Young

University, Provo, UT. 33 pp.

HATTON, S. R. 1932. The fish fauna of Utah Lake. Unpublished master's thesis, Brigham Young University,

Provo, UT. 64 pp.

HAYES, S. P. 1935. A taxonomical, morphological, and distributional study of the Utah Cyprinidae.

Unpublished master's thesis, Brigham Young University, Provo, UT. 115 pp.

HECKMANN, R. A., C. W. THOMPSON, AND D. A. WHITE. 1981. Fishes of Utah Lake. Great Basin

Naturalist Memoirs 5: 107-127.

HOLDEN, P. B., C. N. GOODWIN, AND K. D. THEIS. 1994. A study to determine appropriate water

management actions to enhance native and sportfish populations in Utah Lake: summary of existing

information and preliminary feasibility study. Unpublished report, BIO/WEST, Inc., Logan, UT. 33 pp.

HOLDEN, P., W. WHITE, G. SOMERVILLE, D. DUFF, R. GERVAIS, AND S. GLOSS. 1974. Threatened

fishes of Utah. Proceedings of the Utah Academy of Sciences, Arts, and Letters 51: 46-65.

JORDAN, D. S. 1878a. A catalogue of the fishes of the fresh waters of North America. Bulletin of the United

States Geological and Geographical Survey of the Territories 4(2): 407-442.

1878b. Contributions to North American ichthyology based primarily on the collections of the United

States National Museum. III. B. A synopsis of the family Catostomidae. Bulletin of the United States

National Museum No. 12: 97-237. NATIVE PISCIFAUNA OF UTAH LAKE 73

- --. 1878C. Manual of the vertebrates of the northern United States, including the district east of the

Mississippi River, and north of North Carolina and Tennessee, exclusive of marine species. Second ed.

Jansen, McClurg & Co. 407 pp.

- --. 1880. Notes on certain typical specimens of American fishes in the British Museum and in the Museum

D'HISTOIRE NatureLLE at Paris. Proceedings of the United States National Museum (1879) 2: 218-226.

- --. 1886A. Note on Mr. Garman's paper on The American salmon and trout." Proceedings of the United

States National Museum (1885) 8: 81-83.

- --. 1886B. Identification of the species of CYPRINIDAE and CATOSTOMIDAE, described by Dr. Charles Girard,

in the Proceedings of the Academy of Natural Sciences of Philadelphia for 1856. Proceedings of the

United States National Museum (1885) 8: 118-127.

- --. 1887A. The geographical distribution of fresh-water food-fishes in the several HYDROGRAPHIC basins of

the United States. Pages 133-144 in G. B. Goode, editor, The fisheries and fishery industries of the

United States. Section 3. Government Printing Office, Washington, DC. 178 pp + 32 charts.

- --. 1887B. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic

of Cancer, with notes on the species discovered in 1883 and 1884. United States Commission of Fish and

Fisheries: Report of the Commissioner for 1885 13: 789-973.

---. 1891. Report of explorations in Colorado and Utah during the summer of 1889, with an account of

the fishes found in each of the river basins examined. Bulletin of the United States Fish Commission

(1889) 9: 1-40 + plates 1-5 and map.

---. 1920. The trout of the Rio Grande. COPEIA No. 85: 72-73.

- --. 1928. The distribution of fresh-water fishes. Annual Report of the Board of Regents of the Smithsonian

Institution showing the operations, expenditures, and condition of the institution for the year ending June

30 1927: 355-385 + plate 1.

JORDAN, D. S. AND B. W. EVERMANN. 1896A. The fishes of North and Middle America: a descriptive

catalogue of the species of fish-like vertebrates found in the waters of North America, north of the

Isthmus of Panama. Bulletin of the United States National Museum No. 47, Part 1: 1-1240. GREAT BASIN NATURALIST 74

- --. 1896b. Check-list of the fishes and fish-like vertebrates of North and Middle America. United States

Commission of Fish and Fisheries: Report of the Commissioner for the Year Ending June 30, 1895

21: 207-584.

- --. 1898a. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like

vertebrates found in the waters of North America, north of the Isthmus of Panama. Bulletin of the United

States National Museum No. 47, Part 2: 1241-2182.

- --. 1898b. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like

vertebrates found in the waters of North America, north of the Isthmus of Panama. Bulletin of the United

States National Museum No. 47, Part 3: 2183-3136.

- --. 1900. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like

vertebrates found in the waters of North America, north of the Isthmus of Panama. Bulletin of the United

States National Museum No. 47, Part 4: 3137-3313 + plates 1-392.

- --. 1902. American food and game fishes: a popular account of all the species found in America north

of the equator, with keys for ready identification, life histories and methods of capture. Doubleday, Page

& Co., New York. 573 pp.

JORDAN, D. S., B. W. EVERMANN, AND H. W. CLARK. 1930. Check list of the fishes and fishlike

vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia.

Report of the United States Commissioner of Fisheries for the Fiscal Year 1928 with Appendixes

2: 1-670.

JORDAN, D. S. AND C. H. GILBERT. 1881. Notes on a collection of fishes from Utah Lake. Proceedings

of the United States National Museum (1880) 3: 459-465.

- --. 1882. Synopsis of the fishes of North America. Contributions to North American Ichthyology IV.

Bulletin of the United States National Museum No. 16: 1-1018.

JORDAN, D. S., AND P. L. JOUY. 1882. Check-list of duplicates of fishes from the Pacific coast of North

America, distributed by the Smithsonian Institution in behalf of the United States National Musum, 1881.

Proceedings of the United States National Museum (1881) 4: 1-18. NATIVE PISCIFAUNA OF UTAH LAKE 75

JOUY, P. L. 1882. Description of a new species of Squalius (Squalius aliciae), from Utah Lake. Proceedings

of the United States National Museum (1881) 4: 19.

LA RIVERS, I. 1962. Fishes and fisheries of Nevada. Nevada State Fish and Game Commission. 782 pp.

LA RIVERS, I., AND T. J. TRELEASE. 1952. An annotated check list of the fishes of Nevada. California

Fish and Game 38: 113-123.

LOWDER, J. 1951. A taxonomic study of the Catostomidae of Utah Lake with notes on the fish population.

Unpublished master's thesis, Brigham Young University, Provo, UT. 45 pp.

LOWE, D. W., J. R. MATTHEWS, AND C. J. MOSELEY (editors). 1990. June Sucker: Chasmistes liorus.

Pages 813-814 in The official World Wildlife Fund guide to endangered species of North America.

Vol. 2: pages 561-1180. Beacham Publishing Inc., Washington DC.

MAYDEN, R. L., B. M. BURR, L. M. PAGE, AND R. R. MILLER. 1992. The native freshwater fishes

of North America. Pages 827-863 in R. L. Mayden, editor, Systematics, historical ecology, and North

American freshwater fishes. Stanford University Press, Stanford, CA. 969 pp.

MILLER, R. R. AND G. R. SMITH. 1981. Distribution and evolution of Chasmistes (Pisces: Catostomidae)

in western North america. Occasional Papers of the Museum of Zoology, University of Michigan,

No. 696.

MILNER, J. W. 1874. New species of Argyrosomus and Coregonus. United States Commission of Fish and

Fisheries: Report of the Commissioner for 1872 and 1873 2: 86-89.

MODDE, T., AND N. MUIRHEAD. 1990. Emergence patterns and feeding behavior of larval June Sucker.

Contract 90-0081, Utah Cooperative Fish and Wildlife Research Unit, Department of Fisheries and

Wildlife, Utah State University, Logan. 28 pp.

---. 1994. Spawning chronology and larval emergence of June Sucker (Chasmistes liorus). Great Basin Naturalist 54: 366-370.

MOORE, G. A. 1968. Fishes. Pages 21-165 in W. F. Blair, A. P. Blair, P. Brodkorb, F. R. Cagle, and G. A.

Moore, editors, Vertebrates of the United States. Second ed. McGraw-Hill Book Co. 616 pp. GREAT BASIN NATURALIST 76

OLSON, H. F. 1959. The biology of the Utah Chub, Gila atraria (Girard), of Scofield reservoir, Utah.

Unpublished master's thesis, Utah State University, Logan. 34 pp.

PAGE, L. M., AND B. M. BURR. 1991. Freshwater fishes. Houghton Mifflin Co., Boston. 432 pp. +

48 plates.

RADANT, R. D. AND D. K. SAKAGUCHI. 1981a. Utah Lake fisheries inventory: Executive Summary.

U.S. Department of the Interior, Bureau of Reclamation, and Utah Department of Natural Resources,

Division of Wildlife Resources, Salt Lake City. 33 pp.

---. 198 lb. Utah Lake fisheries inventory. U.S. Department of the Interior, Bureau of Reclamation, and

Utah Department of Natural Resources, Division of Wildlife Resources, Salt Lake City. 244 pp.

RADANT, R. D., AND D. SHIRLEY. 1987. June Sucker: Utah Lake investigations. U.S. Department of

the Interior, Bureau of Reclamation, Contract 8-07-40-S0634, Modification No. 5. Utah Department of

Natural Resources, Division of Wildlife Resources, Salt Lake City. 46 pp.

RADANT, R. D., M. M. WILSON, AND D. SHIRLEY. 1987. June Sucker: Provo River instream flow

analysis. U.S. Department of the Interior, Bureau of Reclamation, Contract 8-07-40-S0634, Modification

No. 4. Utah Department of Natural Resources, Division of Wildlife Resources, Salt Lake City. 45 pp.

RICHARDSON, J. 1836. The fish. Fauna Boreali-Americana; or the Zoology of the Northern Parts of British

America: Containing Descriptions of the Objects of Natural History Collected on the Late Northern Land

Expeditions under Command of Captain Sir John Fran!din, R.N. Part 3: 1-327 + plates 74-97.

RIDE, W. D. L., ET AL. 1985. International code of zoological nomenclature. 3rd ed. International Trust for

Zoological Nomenclature, London. 338 pp.

ROBINS, C. R., R. M. BAILEY, C. E. BOND, J. R. BROOKER, E. A. LACHNER, R. N. LEA, AND

W. B. SCOTT. 1991. Common and scientific names of fishes from the United States and Canada. Fifth

ed. American Fisheries Society Special Publication 20: 1-183.

ROCKWOOD, A. P. 1873. The native fish of Utah. Proceedings of the American Fish Culturists' Association

(1872) 2: 24-25. NATIVE PISCIFAUNA OF UTAH LAKE 77

SCHRENKEISEN, R. 1938. Field book of fresh-water fishes of North America north of Mexico (J. T. Nichols

and F. R. LaMonte, eds.). G. P. Putnam's Sons, New York. 312 pp + 1 plate.

SHIRLEY, D. L. 1983. Spawning ecology and larval development of the June Sucker. Proceedings of the

Bonneville Chapter of the American Fisheries Society 1983: 18-36.

SIBLEY, C. G., AND J. E. AHLQUIST. 1990. Phylogeny and classification of birds. Yale University Press,

New Haven, CT. 976 pp.

SIGLER, W. F., AND R. R. MILLER. 1963. Fishes of Utah. Utah State Department of Fish and Game, Salt

Lake City. 203 pp.

SIGLER, W. F., AND J. W. SIGLER. 1987. Fishes of the Great Basin: A natural history. University of

Nevada Press, Reno. 425 pp. + 8 plates.

SMITH, G. R., AND R. F. STEARLEY. 1989. The classification and scientific names of rainbow and

cutthroat trouts. Fisheries 14: 4-10.

SNYDER, D. E., AND R. T. MUTH. 1988. Description and identification of June, Utah, and Mountain

Sucker larvae and early juveniles. Contribution 37 of the Larval Fish Laboratory, Colorado State

University, Fort Collins. Publication No. 88-8, Utah Department of Natural Resources, Division of

Wildlife Resources, Salt Lake City. 107 pp.

SNYDER, J. 0. 1915. Notes on a collection of fishes made by Dr. Edgar A. Mearns from rivers tributary

to the Gulf of California. Proceedings of the United States National Museum 49: 573-586 + plates 76

and 77.

---. 1922. Notes on some western fluvial fishes described by Charles Girard in 1856. Proceedings of the

United States National Museum 59: 23-28.

STEARLEY, R. F., AND G. R. SMITH. 1993. Phylogeny of the Pacific trouts and salmons (Oncorhynchus)

and genera of the family Salmonidae. Transactions of the American Fisheries Society 122: 1-33.

STUBBS, W. J. 1966. The Weberian osteology of three Utah Suckers (Catostomidae). Unpublished master's

thesis, Brigham Young University, Provo, UT. 53 pp. + literature cited section, tables 1-8, and plates

1-5. GREAT BASIN NATURALIST 78

SUCKLEY, G. 1874. On the North American species of salmon and trout. United States Commission of Fish

and Fisheries: Report of the Commissioner for 1872 and 1873 2: 91-160.

TANNER, V. M. 1936. A study of the fishes of Utah. Utah Academy of Sciences, Arts, and Letters 13:

155-184.

U.S. FISH AND WILDLIFE SERVICE. 1995. Recovery Plan for June Sucker (Chasmistes liorus): Draft. Utah

Department of Natural Resources, Division of Wildlife Resources for Region 6, U.S. Department of the

Interior, Fish and Wildlife Service, Denver, CO. 55 pp.

UTAH LAKE FISH MANAGEMENT ADVISORY TEAM. 1994. Overview of Utah Lake fish management

studies: Draft. Unpublished report. Utah Department of Natural Resources, Division of Wildlife

Resources, Salt Lake City. 52 pp.

UTTER, F. M. AND F. W. ALLENDORF. 1994. Phylogenetic relationships among species of Oncorhynchus:

A consensus view. Conservation Biology 8: 864-867.

YARROW, H. G. 1874. On the Speckled Trout of Utah Lake. United States Commission of Fish and

Fisheries: Report of The Commissioner for 1872 and 1873 2: 363-368.