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Decline of the Shortjaw Cisco in Lake Superior: the Role of Overfishing and Risk of Extinction

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Decline of the Shortjaw Cisco in Lake Superior: the Role of Overfishing and Risk of Extinction Transactions of the American Fisheries Society 139:735–748, 2010 [Article] Ó Copyright by the American Fisheries Society 2010 DOI: 10.1577/T09-019.1 Decline of the Shortjaw Cisco in Lake Superior: the Role of Overfishing and Risk of Extinction CHARLES R. BRONTE* U.S. Fish and Wildlife Service, Green Bay Fish and Wildlife Conservation Office, 2661 Scott Tower Drive, New Franken, Wisconsin 54229, USA MICHAEL H. HOFF U.S. Fish and Wildlife Service, Fisheries Division, 1 Federal Building, Fort Snelling, Minnesota 55111, USA OWEN T. GORMAN U.S. Geological Survey, Great Lakes Science Center, Lake Superior Biological Station, 2800 Lake Shore Drive East, Ashland, Wisconsin 54806, USA WAYNE E. THOGMARTIN U.S. Geological Survey, Upper Midwest Environmental Sciences Center, 2630 Fanta Reed Road, La Crosse, Wisconsin 54603, USA PHILIP J. SCHNEEBERGER Michigan Department of Natural Resources, Marquette Fisheries Research Station, 484 Cherry Creek Road, Marquette, Michigan 49855, USA THOMAS N. TODD U.S. Geological Survey, Great Lakes Science Center, 1451 Green Road, Ann Arbor, Michigan 48105, USA Abstract.—Recent reviews have further documented the decline of the shortjaw cisco Coregonus zenithicus in Lake Superior. This fish was the most abundant deepwater cisco species in Lake Superior in the early 1920s but presently makes up less than 1% of all deepwater ciscoes (i.e., including shortjaw cisco, bloater C. hoyi, and kiyi C. kiyi) captured in biological surveys. Directed overfishing of deepwater cisco species during the 1930s and again during the mid-1960s and 1970s has been suggested as the cause of the shortjaw cisco’s demise. In this paper, we re-examined the overfishing hypothesis by using historical and recent survey data to estimate the proportion of the historical commercial fishery landings that comprised shortjaw ciscoes. We developed time series of estimated harvest and relative abundance for all statistical districts in Michigan waters of Lake Superior during 1929–1996, for which aggregate catch and effort data were available but not previously examined. The spatial distribution of the fishery and the relationships of catch to fishing effort were examined for evidence of overfishing. Our analysis suggested that directed overfishing was probably not the cause of shortjaw cisco demise, as this species appeared to be declining in all statistical districts regardless of the intensity of the fishery. A count-based population viability analysis indicated that quasi-extinction of the shortjaw cisco is highly probable in the near future. We propose an alternative hypothesis based on the decline of Lake Superior’s keystone predator, the lake trout Salvelinus namaycush, which resulted in an expansion of the population of its principal prey, the cisco C. artedi, due to release from predation pressure. Competitive or predation interactions between the cisco and shortjaw cisco may be more likely explanations for the demise of the latter species. Historically, the Great Lakes contained a morpho- were engendered in part by the environment and logically and ecologically diverse assemblage of through hybridization. Many different taxonomic deepwater coregonine fishes (Coregonus spp.), collec- designations were initially proposed, but experts tively known as ‘‘deepwater ciscoes’’ (Koelz 1929). ultimately settled on seven species. Lakes Michigan These fish had highly variable phenotypic features that and Huron contained the most diverse group (bloater C. hoyi, deepwater cisco C. johannae,kiyiC. kiyi, * Corresponding author: [email protected] blackfin cisco C. nigripinnis, shortnose cisco C. Received February 3, 2009; accepted November 12, 2009 reighardi, and shortjaw cisco C. zenithicus), while Published online March 4, 2010 Lake Superior had bloaters, kiyis, and shortjaw ciscoes 735 736 BRONTEETAL. 1989; Todd 2003). Today, Lake Michigan contains only bloaters, Lake Huron has bloaters and very few shortjaw ciscoes, and Lakes Erie and Ontario have lost all of their deepwater cisco species. Lake Superior, in contrast, maintained its original deepwater cisco community, which is composed of bloaters, kiyis, and shortjaw ciscoes (hereafter, the term ‘‘deepwater ciscoes’’ refers to this community rather than to C. johannae; Lawrie and Rahrer 1972, 1973; Bronte et al. 2003). Little genetic differentiation has been found among these fishes, and species designations have been questioned (Reed et al. 1998; Turgeon et al. 1999). The shortjaw cisco, which is the largest of the three deepwater ciscoes in Lake Superior, was once widely distributed throughout the lake and composed more than 90% of the fish landed in the commercial deepwater cisco fishery during the early 1900s (Koeltz 1929; Van Oosten 1937). From the 1920s to 2000s, shortjaw ciscoes declined in abundance lakewide and now make up less than 1% of all deepwater ciscoes (Hoff and Todd 2004; Gorman and Todd 2007). This decline has been attributed to overfishing (Lawrie and Rahrer 1972, 1973; Hoff and Todd 2004; Gorman and Todd 2007) based on trends in reported landings but without any detailed examination of the commercial catch and effort data. Landings of deepwater ciscoes (mostly shortjaw ciscoes and some bloaters) in U.S. waters of Lake Superior were small compared with those in Lake Michigan but were similar to landings in Lake Huron during 1912–1977 (Baldwin et al. 1979; Figure 1). The early Lake Superior fishery in U.S. waters averaged 395,000 kg/year during 1895–1908 (not shown in Figure 1) and then declined and remained low during 1910–1925, with a mean of 38,000 kg/year (Hoff and Todd 2004). Koeltz (1926) reported that deepwater ciscoes, especially shortjaw ciscoes, were ubiquitously abundant but were not in FIGURE 1.—Reported annual harvest of all deepwater cisco demand by the Chicago markets and were fished only species from U.S. waters of Lakes Superior, Michigan, and for local consumption. From 1926 to 1957, harvest Huron during 1915–1977. Data are from Baldwin et al. increased slightly and averaged around 163,000 kg/ (1979). year in U.S. waters, but then increased to an average of 624,000 kg/year during the 1960s and 1970s (Baldwin and Lake Ontario had shortnose ciscoes, bloaters, and et al. 1979), mostly from an expanded fishery in kiyis (Bailey and Smith 1981). Lake Erie had only the Michigan waters (Peck 1977). shortjaw cisco. In addition, the ubiquitous, more Because of its depressed status in Lakes Superior shallow-water cisco C. artedi (formerly known as lake and Huron and its extirpation in Lakes Michigan and herring) was found in all of the Great Lakes. Ontario, the shortjaw cisco is being considered for Overfishing, introduction of exotic species that altered listing under the Endangered Species Act of 1973 in the fish communities, and hybridization with closely the United States and was recommended for listing as related species during the last century have been ‘‘threatened’’ by the Committee on the Status of proposed to explain the extinction of most forms Endangered Wildlife in Canada (Todd 2003), but this (Smith 1964, 1968, 1970; Berst and Spangler 1972; recommendation has not yet been acted upon. A Christie 1972, 1974; Wells and McLain 1972; Hartman recovery plan for the species is in preparation for 1973; Scott and Crossman 1973; Todd and Stedman Canadian waters, while no actions are being considered SHORTJAW CISCO DECLINE IN LAKE SUPERIOR 737 FIGURE 2.—Map of fishery statistical districts (MS-1–6) in the Michigan waters of Lake Superior (Hile 1962). for U.S. waters pending a listing decision by the U.S. the Michigan Department of Natural Resources Fish and Wildlife Service (USFWS). thereafter. Pursuant to this situation, we re-examined the Catches of the cisco (i.e., lake herring) were reported historical information on the 1929–1996 commercial separately from deepwater cisco catches, but unfortu- fishery for deepwater ciscoes in the Michigan waters of nately the design of the report forced all deepwater Lake Superior as well as observations from Koeltz cisco catches to be grouped under a single column (1929) in the 1920s and fishery-independent survey entitled ‘‘chubs, including blackfins’’ (referring to the data for 1953–2004 from the USFWS and U.S. vernacular for deepwater ciscoes). Hence, we needed to Geological Survey (USGS), as previously published partition shortjaw cisco catches from the total reported by Hoff and Todd (2004) and Gorman and Todd landings. Recent reviews of trends in shortjaw ciscoes (2007). Our objectives were to (1) develop the first from the 1920s through 2004 (Hoff and Todd 2004; time series of relative abundance of shortjaw ciscoes in Gorman and Todd 2007) provide point estimates (years Michigan waters of Lake Superior from 1929 to 1998, where data were available) of the proportion of (2) re-examine the shortjaw cisco decline relative to the shortjaw ciscoes and bloaters captured in small-mesh ‘‘overfishing hypothesis’’ in light of historical catch and gill-net assessments in Michigan waters of Lake effort data not considered previously, and (3) provide Superior. These estimates were taken from assessment the first estimates of the probability of extinction using gill nets set at depths of 40–160 m; the assessment gill the new relative abundance data from commercial nets had 64–70-mm stretch-measure mesh (1922, catches. Furthermore, as a result of our analysis of 2000–2004) or 51–76-mm mesh (1953, 1958–1992). objective 2, we developed a new hypothesis for the The 1922 studies by Koeltz (1929) used mesh sizes that decline of the shortjaw cisco in Lake Superior. ranged from 54 to 76 mm although meshes of 64–70 mm were most common. These mesh sizes were similar Methods to those used in the commercial fishery, which mostly Time series development.—We developed a time ranged from 64 to 70 mm, although smaller and larger series of the relative abundance of shortjaw ciscoes meshes were occasionally used (Koeltz 1929; Ander- from the commercial small-mesh gill-net fishery in the son and Smith 1971).
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