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Ecology of Upper Klamath Lake Shortnose and Lost River Suckers

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Ecology of Upper Klamath Lake Shortnose and Lost River Suckers ECOLOGY OF UPPER KLAMATH LAKE SHORTNOSE AND LOST RIVER SUCKERS 4. The Klamath Basin sucker species complex 1999 ANNUAL REPORT (partial) SUBMITTED TO U. S. Biological Resources Division US Geological Survey 104 Nash Hall Oregon State University Corvallis, Oregon 97331-3803 & Klamath Project U. S. Bureau of Reclamation 6600 Washburn Way Klamath Falls, OR 97603 by Douglas F. ~arkle', Martin R. ~avalluzzi~,Thomas E. owli in^^ and David .Simon1 1Oregon Cooperative Research Unit 104 Nash Hall Department of Fisheries and Wildlife Oregon State University Corvallis, Oregon 97331-3803 E -mai1 : douglas.markle@,orst.edu 2Department of Biology Arizona State University Tempe, AZ 85287-1501 Phone: 480-965-1626 Fax: 480-965-2519 E -mai 1 : [email protected] July 26, 2000 There are 13 genera and 68 species of catostomids (Nelson 1994) with three genera and four species occurring in Klamath Basin (Bond 1994)- Catostomus rimiculus Gilbert and Snyder, 1898 (Klamath smallscale sucker, KSS), C. snyderi Gilbert 1898 (Klamath largescale sucker, KLS), Chasmistes brevirostris Cope, 1879 (shortnose sucker, SNS), and Deltistes luxatus (Cope, 1879) (Lost River sucker, LRS). Lost River and shortnose suckers are federally listed endangered species (U.S. Fish and Wildlife Service 1988). The four Klamath Basin suckers are similar in overall body shape, but highly variable, and are distinguished by feeding-related structures, adult habitat and geography. The two Catostomus species have large lips, widely-spaced gillrakers, and are primarily river dwellers with C. snyderi mostly found in the upper basin and C. rimiculus in the lower basin and adjacent Rogue River. Deltistes luxatus has smaller lips, short "deltoid" Catostomus-like gillrakers, and is primariliy a lake dweller. Chasmistes brevirostris has small lips, many closely-spaced gillrakers with secondary branching, and is also primarily a lake dweller (Andreasen 1975, Miller and Smith 1981, Buettner and Scoppettone 1991). Catostomids were among the first freshwater fish known to hybridize in nature (Hubbs, et al., 1943). Miller and Smith (1981) stated they "had not seen any recently- collected specimens from (Upper) Klamath Lake that are the same as brevirostris", that the traits indicated introgression with C. snyderi, and that 'none of the available names is applicable" to Klamath Basin populations. This and other problems with species identification prompted a morphological and genetic study of Klamath Basin suckers. The objectives were to understand sources of variation, provide field biologists with usable identification criteria, and provide initial identifications for a multi-investigator sucker genetics program, referred to herein as KLAMGEN. STUDYAREA The KLAMGEN specimens were collected from the Klamath and Rogue river basins in south central Oregon and northern California (Fig. 1). Klamath Basin collections targetted five subbasins (Fig. 1). The Upper Williamson subbasin is that part of the Williamson R. above Klamath Marsh. The Sprague subbasin is the Sprague R. above Chiloquin Dam including the Sycan R. Specimens caught in the ladder at Chiloquin Dam were considered to be in the Sprague subbasin. Upper Klamath subbasin includes Upper Klamath Lake, the lower Williamson R., lower Sprague R. downstream of Chiloquin Dam, and the Link River downstream of Klamath Falls. The Lost River subbasin includes Clear Lake, Lower Klamath Lake and Gerber Reservoir. The Lower Klamath subbasin includes the three downstream reservoirs (J. C. Boyle, Copco, and Iron Gate) and Jenny Cr. Specimens collected outside the KLAMGEN program were also assigned to the appropriate subbasin. See U.S. Department of the Interior, Bureau of Land Management (1995) for detailed descriptions of the climate, geology, and topology of the Klamath Basin. MATERIALSAND METHODS Abbreviations for museum acronyms follow Leviton -et -al. (1985). We examined 1782 suckers - 1741 from the Klamath Basin and 41 from the Rogue River. The most extensive data collection was on 333 adult specimens collected for the KLAMGEN project (Table I), of which 325 carcasses were deposited in the Oregon State University Fish Collection (0s). Eight specimens were sampled non- lethally for tissues. The KLAMGEN protocol attempted to obtain samples from suspected spawning groups in spring and early summer, 1993-1994, but 183 (55%) were collected outside the spawning seasons in August-November 1993. Much additional Klamath material (1085 specimens) was obtained from juvenile collections (Markle and Simon unpublished), also deposited at OS and radiographs of the holotype of C. brevirostrisi (ANSP 20950), holotype of Ch. stomias (USNM 48223, and holotype of Ch. copei USNM 48224. Unless noted all analyses are based on the entire data set. Abbreviations and descriptions of counts and measurements are found in Table 2 and generally follow Hubbs and Lagler (1964). All vertebral and vertical fin ray counts were taken from radiographs and do not include the four Weberian centra. All measurements were in mm. Lip area and perimeter were calculated for 32 specimens using a digital imagery system equipped with Optimas 5.0 (1995) software. Not all variables were measured for all specimens because of damage or lack of variation detected after preliminary analyses. Data were analyzed using STATGRAPHICS Plus (1994- 1996). Univariate characters were evaluated using Kruskal- Wallis non-parametric ANOVA to test for significant differences with comparisons adjusted using the Bonferroni inequality. Multivariate analyses used Principle Components Analyses (PCA) to reduce data and uncover data structure and Discriminant Functions Analysis (DFA) to classify individuals. DNA was extracted from 324 KLAMGEN individuals and initially screened with several sets of primers from the cytochrome b and ND4L genes. Those that amplify the ND4L gene were selected because of ease of scoring and sequencing. Sixteen haplotypes were identified from 324 individuals (Table 1) . RESULTS Lip morphology and initial identification. ---Initial field identifications of C. snyderi and Ch. brevirostris were inconsistent. In Upper Klamath, Sprague, and Upper Williamson subbasins both species were identified based on overall appearance, primarily lip morphology, whereas a single species (Ch. brevirostris) was identified in the field in Lost River subbasin. We surveyed our quantitative and categorical lip morphology characters (below) and re- identified Lost River Ch. brevirostris as C. snyderi if the following combination of characters was present in an individual: 1) symphyseal lower lip gap absent, teratological, or present anteriorly but lips touch posteriorly (LLGAP= N, T or A) , and 2) length of contact of lower lip lobes greater than 50% of eye diameter (GAPLMM/EYE>o.5) . All subsequent analyses are based on these identifications and all descriptions of lip morphology are based on KLAMGEN specimens. Lips of Catostomus species were larger with more surface area relative to the lip perimeter than lips of D. luxatus or Ch. brevirostris (Fig. 2) . Based on a Bonferroni multiple range test, the means of the AOMMM/POMMM ratio sorted into two significantly different homogenous groups, Catostomus species and D. luxatus plus Ch. brevirostris. The position of the posterior margin of the lower lip (LLLRM) was scored as anteriad, even or posteriad of the ventroposterior corner of the maxilla (Fig. 3). Specimens with small lips ending anteriad of the maxilla also had a short distance between the symphysis of lower jaw and the point where lower lip lobes separate (GAPLMM) (Fig. 4). Specimens with larger lips extending posteriad of the maxilla had larger GAPLMM while those with lips even with the maxilla were bimodal for this character (Fig. 4). Expressed as a ratio to eye diameter, the GAPLMM measurement fell into two categories, less than or greater than 50% (Fig. 4). Lips were posteriad of the end of the maxilla in 100% of C. rimiculus, even or posteriad of the maxilla in 98.9% of C. snyderi, even or anteriad of the maxilla in 95.6% of Ch. brevirostris, and anteriad of the maxilla in 100% of D. luxatus. The presence or absence of a symphyseal gap between the lobes of the lower lip (Fig. 3) was also related to the GAPLMM/EYE ratio in specimens (Fig. 5). When a gap is present, the ratio is smaller. Again, the GAPLMM measurement fell into two categories, less than or greater than 50% of the eye diameter (Fig. 5). Catostomus species generally had no lower lip gap, but 9.1% of C. rimiculus (n=55) and 8.4% of C. snyderi (n=95) had lower lip gaps. In Ch. brevirostris, 100% of Sprague and Upper Klamath specimens possessed a lower lip gap and 72.7% of Lost River specimens had a lower lip gap. All specimens of D. luxatus had a lower lip gap. The lower lip gap developes early and is present in young-of-the year juveniles (Fig. 6). In 103 Ch. brevirostris 27.9-90.2 mm SL from Upper Klamath Lake (0s 13969 & 13982), two lacked a lower lip gap while all 51 C. snyderi from the Upper Williamson R. (0s 13882) lacked a lower lip gap. Lip and lower jaw deformities complicate interpretation of these structures. Two Upper Williamson C. snyderi had incomplete development of the branchiostegal membranes creating a long gap extending from the isthmus to the symphysis of the lower jaw (0s 015903-A, B). Other specimens had distorted jaws and lips which we attributed to fixation, but which could be deformities. DNA.---Most haplotypes (n=10) were rare, occurring in only one or two individuals, while the remaining six haplotypes were found in at least 12 individuals (Table 3, Fig. 7) . Individuals of C. rimiculus always exhibited haplotypes 'I" in Klamath Basin and haplotype 'L" in the Rogue R. and only 'I" was found in other species. Each C. rimiculus haplotype was more similar to the common haplotype of D. luxatus than to each other. Most D. luxatus (33 of 40) had haplotype "K" that was rarely found in other species. Haplotype 'N", that appears to be derived from haplotype 'K", was more common in Ch.
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