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Fr DC 02 cm-4. Ant-60,1cl i'arks &RC 1.44r•;.-, L t.hztic St( wit viNs C Pr4La4s j s r , 5. t 3-1 .t1) . 6114 , -1 SPECIES RELATIONSHIPS AMONG FISHES OF THE GENUS1 GILA IN THE UPPER COLORADO RIVER DRAINAGE 2 2 3 Gerald R. Smith , Robert Rush Miller , and W. Daniel Sable Taxonomic treatment of the chubs of the Gila hump, and coded continuous morphometric variables robusta complex (family Cyprinidae) that inhabit (23.1 15.8, etc.) into discontinuous states ("A", the Colorado River basin and certain rivers in "B", etc.), thus losing much of the resolving north-western Mexico (Figure 1) is problematical power of the measurements. The clustering illus- and requires study of the entire group for final trated by them shows only the level at which spec- resolution. It is intended to do this, but in this imens join clusters; it does not show intermedi- paper we present data demonstrating that there are acy, and it cannot display the morphological re- three species of this complex in the Colorado lationship of a specimen between other specimens. River and its major tributaries, from the Grand Thus there is no way to evaluate the supposed Canyon region upstream. These chubs are the most "intergrades" cited by Holden and Stalnaker: in extreme members of the genus in their specializa- their paper the "intergrades" may appear as a tion for life in the unique big-river habitat of result of the method of coding data. They were the Colorado River. unable to determine whether the so-called inter- grades were part of one variable complex or two Our objectives are (1) to determine the valid- incompletely separated species. In contrast, we ity and relationships of the roundtail, bonytail, find that there are very few intermediates, in and humpback chubs in the big-river habitat; the Green River at least, and that most specimens, (2) to determine their relationships to other including the "intergrades" shown by Holden and populations of Gila in the middle and upper Stalnaker (1970: Figure 4), can be assigned to Colorado River basin; and (3) to find characters one of the two species cypha or elegans. useful for identifying young and adult Gila robusta Baird and Girard, G. elegans Baird and Our analysis consists of two parts. First, Girard, and G. cypha Miller, and possible hybrids. principal components analysis of 34 meristic and We do not at this time treat the complex repre- morphometric characters, not including a direct sentatives of Gila in the Gila River basin (see measure of the nuchal hump, is used to establish Rinne, 1976), except to note that G. intermedia clear evidence of morphological segregation of (Girard) may be a separate evolutionary line, the large-river specimens into three distinct secondarily related to G. robusta. clusters. It is important to establish that the segregation is not based on the nuchal hump or Misinterpretation of the patterns of variation other subjective characters, but on a broad range in the nuchal hump and lack of decisive, objective of body proportions, fin-ray counts, and verte- evidence has been the main cause of disagreement bral numbers, presumably with a broad genetic among those studying the G. robusta complex. The basis. Second, after establishing the existence nuchal convexity has never been measured precise- of three separate populations, with little or no ly or quantitatively related to other characters. overlap, at least within the unmodified, large- For example, Holden and Stalnaker (1970) used river habitat, individual discriminating key char- subjective code values for the form of the nuchal acters are developed for identification of these fishes in the laboratory and field. Such charac- 1 ters are critically needed to end the confusion Completion of this paper was delayed by clo- that has clouded attempts to formulate a sound sure of upper Green River to scientists other than management policy for these endangered or threa- federally funded personnel during the five-year tened fishes. The population analysis and the period 1963-1967. Considerable aid was received development of key characters were conducted from a number of organizations and individuals. separately by us in order to avoid circular taxo- Included are research grants to R. R. Miller from nomic logic and to enable the two systems to serve the Horace H. Rackham School of Graduate Studies as useful tests of one another. (1950), the National Science Foundation (G-15914, 24129, 24465, GB-3271, 4854, 6272X), and the Na- Principal components analysis, a multivariate tional Park Service (1975), with cooperation from statistical method enabling taxonomic use of com- the various western states for collecting permits. bined (correlated) information from many charac- Frances H. Miller recorded and calculated data, ters, was applied to 34 characters of 140 speci- helped prepare the distribution map, and criti- mens. These characters are: dorsal, anal, pec- cized drafts of the manuscript. Jeffrey N. toral, and pelvic fin-rays (rays from both paired Taylor prepared the original plots for Figure 1. fins recorded); standard length, head length, eye Loans of specimens were received from the U.S. diameter, snout length, preanal length, head National Museum (E. A. Lachner, S. H. Weitzman, depth through eye, head depth at occiput, inter- W. R. Taylor, Susan Karnella), Arizona State Uni- orbital width, occiput to tip of snout, dorsal- versity (W. L. Minckley), Bell Museum of Natural fin basal length, anal-fin basal length, predorsal History (Samuel Eddy, H. B. Tordoff, C. W. Huver), length, pectoral-fin length, pelvic-fin length, and the Museum of Northern Arizona (S. E. Caroth- upper jaw length, mouth width, body depth over ers). Mark J. Orsen (of the Museum of Zoology) pelvic insertion, caudal-peduncle depth, anal ' and Karna M. Steelquist (of the Museum of Paleon- origin to caudal base; number of vertebrae; pharyn- tology) prepared the drawings and other illustra- geal arches, total length, width, length of tions, except Figure 10, taken by Bruce J. Turner. anterior limb, and length of posterior limb, of Abbreviations are: UMMZ (University of Michigan both left and right arches (eight characters). Museum of Zoology) and USNM (U.S. National Museum This analysis discriminated the populations in of Natural History). question (Figures 2-5) showing the major trends 2 in variation, the characters dominant in the Museum of Zoology, University of Michigan, trends, and the characteristics of typical as well Ann Arbor, MI 48109. as intermediate individuals. 3 Wisconsin State University, Whitewater, A graph of 140 individuals plotted according to Wisc. 53190. their scores on principal components I and II, 613 121 117 113 25 0 25 50 100 200 SCALE OF MILES 50 0 50 100 200 300 SCALE OF KILOMETERS 1 - Gila River • Gila reboto 2 - Little Colorado River O Gila cyplso 3 - Colorado River O Gila elegans I • Son Juan River 5 - Green River 117 113 109 105 FIGURE 1. Distribution of Gila robusta, Gila elegans, and Gila cypha. See Rii-i-fteRAPme (1976) for distribution of allied forms in the Gila basin. 614 • 76 % elegans U. 9% 00 00 o o 00 0 o 0 0 0 0 0 0 q0 cypha 0 0 0 0 0 0 / 0 o 0 0 O 0 ® 0 CE1 En 0 0 00 O DOD 0 ® 0 0 O DD 0 _04* DO 0 00 0 • • 0 0 0 0 0 • • O 00 ® • e 8 •• ® *seminude ® ••• • .• .:• ----- 0 • % ib • *io • • g • • • - • • robusta -----------..,....„.. ....,......... •• FIGURE 2. Principal components analysis, components I and II, of four forms of Gila. G. robusta robusta, G. elegans, and G. Cypha are discriminated by the combination of projections shown in Figure 2 and 3. G. r. seminuda is scat- tered through the other clusters. Percentages indicate the proportion of the total variation accounted for by each component. TABLE 1. Development of nuchal hump in Colorado River chubs, Genus clia.a Species Ratio (range) S.L.range (mm) cypha (31) 6-13 206-328 elegans (20) 15-29 245-412 r. robusta (17) 28-207 211-309 r. seminuda (4) 31-121 175-247 a — Expressed as ratio: Pi rn I '-'2 distance Depth frontal depression Figures in parentheses indicate number of speci- mens measured. 615 the Colorado and Green rivers. This conclusion for the necessary stability to persist. cannot be confidently applied to certain popula- tions that we have available but have not fully In a period of controversy following the poi- studied from Lake Powell, however. Furthermore, soning of the upper Green River in 1962 (Miller populations of Gila robusta from certain tribu- 1963b), erroneous statements about the classifi- h/ taries ,ere elegans and cypha are absent parallel cation, biology, and distribution of Colorado these species in many characteristics. We choose River chubs appeared which have been destructive a taxonomic treatment of this situation that em- to the understanding and management of these phasizes the specific distinction of the three fishes, and require correction. For example, populations in the big-river habitat, but admit Stroud (1963:7) irresponsibly claimed that the the possibility that the species isolating mecha- construction of Flaming Gorge Reservoir resulted nisms may break down under disturbed (reservoir) in making available "large numbers" of humpback conditions and that the populations in tributary chubs, all of which were supposedly males. The streams may not be completely independent or iso- agency (U.S. Bureau of Sport Fisheries and Wild- lated from the three central forms. We refer to life) that "confirmed the presence and ready the entire complex as the Gila robusta super- availability of numerous humpback chubs in [the species, including robusta, cypha, and elegans Green River below Flaming Gorge Dam] . ." as defined above, and including subspecies or subsequently reported (Vanicek 1967; Vanicek et races of robusta, seminuda of the Virgin River, a al.