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Home , Bluegill, Centrarchidae, Lepomis, Pumpkinseed

Scientific Name (Linnaeus): gibbosus

Stephannie Spurr

Fish 497

Fall 2008

Diagnostic Information native range, reach an average : length of 127-190 mm. In their Non-native Family: , habitats they are smaller, on average 80-100mm. :Lepomis This has been attributed to a change in resource : gibbosus allocation from growth to reproduction in order Common Names: sunfish, yellow to increase their establishment in a novel sunfish, common sunfish, sunfish, round environment. Establishment in non-native range sunfish, punky, sunny, sun , pond perch, has also been successful as many introduced bream (Becker 1983) areas lack a native predator such as bass, and where bass is present, the native fish are a Basic Identification: Deep, laterally preferred prey rather than pumpkinseed. compressed body. Average length 127-190 mm. Phenotypic plasticity is also seen in their non- Short terminal mouth. Short, stubby gillrakers; native range in body shape allowing them to almost as wide as they are long. Olive colored exploit a larger range of habitat. Pumpkinseed sides with orange to red-orange belly and breast. fish have been widely stocked in the Pacific Blue and orange cheek stripes, and a red to Northwest for sportfishing by the WDFW. orange spot located posterior of the opercular Management of unauthorized stocking is flap. There are two dorsal fins, but they appear through the use of rotenone. The WSANS places as a single fin. Anal fin with spines, the pelvic pumpkinseed on their “Regulated Species” list fin is located thoracic, and the pectoral fin is where as they are on the WDFW’s prohibited long and when laid for ward towards the eye, list. reaches the front of the eye. The caudal fin is slightly forked. (Wydoski and Whitney 1979; Origin and distribution. Becker 1983). During breeding season, males will be more brightly colored and the females History of invasiveness. will have darker more prominent vertical bands (Wydoski and Whitney 1979). The first introduction of pumpkinseed into the Pacific Northwest is believed to have occurred in the of in Overview 1893 (Wydoski and Whitney 1979). The U.S. Pumpkinseed fish are a Fish and Wildlife Commission released in the family Centrarchidae. Pumpkinseed are and a small sunfish fry of native to the Great Lakes area, but have been which pumpkinseed was presumed to be globally introduced for sportfishing. In their included. Wide spread stocking has occurred in Figure 1: Native and Non-native Distribution through-out the (United States Geological Survey 2007). over 160 lakes (Washington escaped into the wild. Pumpkinseed Department of Fish and Wildlife. 2005), for pan introductions and naturalizations continued to fish fisheries, promoting a juvenile pan fish occur through out in 35 countries derby, (Sportsman Lake Stock Assessment including Czechoslovakia, Italy, Poland, Survey 2001) and for general sport fishing.( Switzerland, Hungary, Romania, the former Department of Fish and Wildlife. 2005). Sunfish Yugoslavia, The Netherlands, and Belgium. spread into British Columbia is most likely the (Raveret-Wattel 1900; Welcomme result of its natural spread from the Columbia 1981;Grimaldi 1972;Muus and Dahlstrøm 1968; River system (Crossman 1991). Outside of the Thienemann 1950; Nikolsky 1961;Looyen 1948; United States, pumpkinseeds were imported to Brouwer 1925; De Groot 1985) Pumpkinseeds by Max Von dem Borne in an are also wide spread in the Iberian Peninsula experimental attempt to raise them in his (García-Berthou and Moreno-Amich 2000) and hatchery. In 1885 pumpkinseeds arrived in parts of Southern England (Künstler 1908). France and were subsequently released, or

Current distribution in the Pacific Northwest Pumpkinseed are presently found in over 160 lakes in Washington state, 11 documented lakes in Idaho and 66 lakes in Oregon (Washington Department of Fish and Wildlife. 2005; Simpson and Wallace 1978; Lee et al. 1980; Lamp man 1946; Chapman 1942). It is also found in British Columbia in the Columbia River (Crossman 1991).

Life-History and Basic Ecology

Life Cycle Figure 2:. Growth rate curve compiled from pooled Life for a pumpkinseed starts off with a data and fitted with von Bertalanffy growth model 1mm diameter egg stuck to a small stick or rock for European and North American pumpkinseed (Copp et al. 2004). inside an under water nest. Hatch time is temperature dependant with a range of 47 hours macroinvertibrates to gastropods. This change at temperatures between 19.0-24.7° C and 3 has been correlated with rapid growth in their days at 27.8°C (Balon 1959). The larval stage native environment. Secondly, the European lasts 22 days. (Houde and Zastrow 1993). Young pumpkinseed shows an in increase in resource pumpkinseed leave their nest but remain near the allocation to reproduction, rather than growth. shallow breeding area (Becker 1983) and have They state that a higher reproductive effort been observed to travel in loose schools (Emery increases the chance of establishment in a novel 1973). Growth rates can vary widely depending environment. In their native range pumpkinseed on location. are sexually mature by 2 to 3 years of age, can In a study done by Copp et al. (2004), growth live to 12 years, but generally not beyond 6 rate and adult size was lower for introduced years (Wydoski and Whitney 1979). In their pumpkinseed in Europe than pumpkinseed in its introduced ranges, ages of sexual maturity vary. native range in (figure 2). They In Southern England, pumpkinseeds are not propose this is due to two factors. First, unlike found to be mature until almost 4 years (The the adult native pumpkinseed, the European Fisheries Society of the British Isles 2006.). In adult pumpkinseed have no dietary transition France, reproductive maturity was achieved in from the juvenile food preference of soft-bodies the summer after their first winter and the life span in this reservoir is 3years (Dembski et al. The dietary shift is not regularly seen in Europe. 2006). Both these exceptions were related to Lack of gastropods led to less trophic temperature difference from the native range. In segregation between adults and juveniles and France the adverse biotic condition of high water has been proposed as one mechanism to explain temperature resulted in high adult mortality and decreased growth, as well as diminished resulted in early, highly fecund reproduction, predatory pressure (Copp et al 2004) and when water was colder in southern England, Molluscivory is seen at Lake Banyoles on the development time was slowed. There is also the Iberian Peninsula. Endemic are consumed lack or absence of predators, in the non-native by adults as well as and . environment, which can result in delayed Juvenile fish fed on littoral microcrustaceans. maturation as a fish grows larger (Reznick and (Garcia-Berthou & Moreno-Amich, 2000). This Endler, 1982). has led to life history traits and morphology changes in native snails (DeWitt, 1998). Feeding Habits Negative impacts on the snails have been seen as In the native habitat, pumpkinseeds feed a potential problem (Garcia-Berthou & Moreno- on different species depending on their life Amich, 2000). stage. Juvenile pumpkinseed feed on soft-bodied The pumpkinseed is active during the day, but macroinvertibrates (Copp et al. 2004). Adult by dusk they move to the bottom to rest near pumpkinseeds are omnivorous and will feed on fallen logs or rocks and rest (Becker 1983). several prey species including , worms, During winter time, the pumpkinseed doesn’t insect larva and plant material. Unique to adult feed, and the stomach is shrunken and filled with pumpkinseeds in relationship to other sunfish is mucus. Once the water begins to warm, a its specialized large jaw musculature with large feeding response is initiated (Carlander 1977). molar like teeth that allows it to feed on snails (Becker 1983; Robinson et al. 2000). The shift Reproductive Strategies in food type with more emphasis on Pumpkinseeds are iteroparous, thus they molluscivory parallels sexual maturity, but is not several times within their life span. evidence of a preference for snails, but just of Spawning in native regions occurs in early May their availability, the morphological structured to August (Becker 1983). Temperatures between jaw to bite through the shells, as well as an 11.5 and 14.0 C are needed to trigger male’s ontogenetic shift in food resource that allows gonad growth, and temperatures between 14.0 greater partitioning between the adults and the and 16.5 C are needed are need to initiate female juveniles (Garcia-Berthou and Moreno-Amich, gonad growth (Burns 1976). The larger males 2000). construct a nest which is a circular depression on the substrate of the warmer, shallow, littoral have two nests at the same time (Breder and zone. This is done by fanning the tale and Rosen 1966). removing large objects with their mouths Males guard the nest from potential threats, (Adams and Hankinson 1926). Once the nest is including sneaker males, because cannibalism built, males will defend their territory around the occurs in pumpkinseed (Rios-Cardenas nest from about a 1 meter in diameter inwards. and Webster 2005). They also fan the eggs Defense of the nest includes nipping, chasing, which also results in higher survival rates but fanning out the opercula, and very rarely actual this parental care costs the males body mass as mouth to mouth fighting (Clark and Keenleyside they don’t leave the nest to feed (Rios-Cardenas 1967). Small, sexually mature males don’t make and Webster 2005), ultimately suffering high nests, rather these “sneaker” males insinuate mortality rates by the end of the season (Gross themselves into the spawn of the males who 1980). Paternal guard over the nest usually last built the nest, and the female he has attracted about nine days (Carlander, 1977) but can drop- (Gross 1979). With the intrusion of a sneaker off at the wriggler stage on day four when the male, the parental males looses some measure of males begin to clean their nests for the next paternity, and expends energy raising young that spawn (Rios-Cardenas 2003). offer no genetic advantage to his lineage (Rios- Environmental Optima and Tolerances Cardenas and Webster 2005). Courtship behavior begins when the male chases In the Pacific Northwest, pumpkinseed a female who has come close to his nest. The fish thrive in lakes that have thick vegetation. female swims back to an aggregate of females in Clear slow moving water is preferred, but in deeper waters, and this happens multiple times waters without aquatic weeds, their numbers are before the female submits to the male’s much lower. They can be found in slow-moving directional drive into his nest. Male and female rivers as well as lakes. Summer water swim next to each other , move in a circle of temperatures of over 70° are preferred. Slow around 11 circuits a minute. During this time the growth and relative small size in comparison to female rolls to her side and while she releases fish growing in their native size has been eggs, the male release sperm (Becker 1983). attributed to cooler Pacific Northwest waters, This needs to happen simultaneously as eggs are and this has led to decease in their desirability as only viable for less than a minute (Bagenal and a sport fish (Wydoski and Whitney 1979). Braum 1971).Females produce an average of Biotic associations 2,436 eggs (Wydoski and Whitney 1979).Over The pumpkinseed fish is parasitized by the next two days the male will be receptive to several species of the cnidarian myxosporidan. more females (Gross, 1980). Males may also They are found in their eyes as well as their Invasion Process pancreas, ureters and urinary bladder. This is the same family of microscopic cnidarian parasites Primary pathway of introduction into the that are responsible for whirling disease in Pacific Northwest of pumpkinseed was stocking salmonids. (Cone and Anderson 1977; Minchew for recreation. (Washington Department of Fish and Sleight 1977). They are also host to two and Wildlife. 2005), Further spread has been freshwater mussel glochidia, Amblema plicata attributed to the natural dispersal from the and Andonta implicate (Hart and Fuller, 1974). Columbia River system. Pathways for European Pumpkinseed is sympatric with another sunfish, introductions are more diverse. García-Berthou the , in the North American Midwest. et al. (2005) have stated that European invasions There is some competition between the two are motivated by more sociological and species for habitat, and food, but is largely historical factors, rather than ecological ones. avoided with the morphological plasticity They illustrate this with the example of the capabilities of the pumpkinseed fish. When in largemouth bass, salmoides, (a common waters, the adult pumpkinseed will species found with pumpkinseeds in their native inhabit the littoral zone, and feed on benthic range) and their successfully establishment in invertebrates, while the adult bluegill inhabits many African countries. The conclusion was that the limnetic zone and feeds upon if the bass invasion was due to a natural (Werner and Hall 1976). In lakes without dispersal, pumpkinseed would have spread with bluegill, adult pumpkinseeds are found in the them, when in fact, they haven’t. limnetic zone as well as the littoral zone Pumpkinseed introduction into southern England (Robinson et al, 2000). has initially been attributed to pond stocking for During the juvenile stage for pumpkinseed and ornamental reasons. (Copp et al. 2002). The bluegill sunfish, there is more competitive spread has been attributed to two factors; one, pressure on the pumpkinseed. Both sunfish’s the downstream movement from lakes connected offspring inhabit the littoral zone. Lakes have a to streams, and two, hitchhikers on intentional higher proportion of zooplankton, which is the fish transfers. Small pumpkinseeds were seen adult bluegill’s food, versus benthic under the opercula of large cyprinids during invertebrates such as snails that are adult their transfer (Villeneuve et al. 2005). pumpkinseed prey. With a better adult dietary Pumpkinseeds have been fairly restrictive in resource, the bluegill contributes more offspring their spread of England. Limited food supplies, to the shared littoral zone decreasing resource less than ideal climate and lack of establishment availability for the young pumpkinseed in streams where they have been introduced (Mittelbach and Chesson 1987) have been offered as explanations (Klaar et al. 2004). Some of these factors could be overcome in the breeding season with no dependence on in the future. Propagule pressure in the form of size. This adaptation was attributed to a change continued introduction of fish from the main in energy expenditures towards reproduction source to the out lying streams could lead to rather than tissue growth due to early adult successful establishment, as could a climate mortality from the increase in temperature change due to global warming. (Klaar et al. (Dembski et al. 2006). 2004; Villeneuve et al. 2005). Opportunistic life strategies are an important In their natural habitat pumpkinseeds make up aspect of the pumpkinseeds ability to establish in part of several predator fishes’ diet, including novel territories, and this is most evident in the the largemouth bass Micropterus salmoides Iberian Peninsula. Pumpkinseed fish were (Becker 1983). With the exception of a few accidentally introduced by the Catalonia discrete locations within Europe, and the Iberian Regional Fisheries Service while stocking Peninsula, largemouth bass were not included in largemouth bass for anglers (Elvira 1995). There the introduction of pumpkinseed (Keith & are no native piscivorous fish on the Iberian Allardi 2001). One naturally occurring fish Peninsula. on pumpkinseed by bass, predator, the pike-perch Sander lucioperca, which occurs in their native range, is not as feeds only sparingly on pumpkinseed (Wheeler prevalent on the Iberian Peninsula. One 1968). Lack of predation is thought to a reason explanation that has been presented is the for the smaller size of the non-native abundance of fish with slender bodies (versus pumpkinseed in relationship to native the wider pumpkinseed body) in comparison to pumpkinseeds due to lack of selective pressure North America in which the largemouth bass can to have a large body in order to escape the gape prey upon (Bhagat et al. 2006). size of predatory fish such as bass (Copp et al Ecological impact from pumpkinseeds has been 2004). This switch in size contributed to a widely studied in Iberia and their threat to native change in the pumpkinseed fish being a mollusks, predation on fish eggs, piscivory and desirable market fish to a nuisance fish in France direct competition for food identified as causes (Roule 1931). Another life history characteristic for declines of native fauna (García de Jalón et that was observed to have been altered in Europe al., 1993; García-Berthou & Moreno-Amich was recorded in the Mirgenbach reservoir of 2000; Godinho & Ferreira 1997). France. Mirgenbach reservoir has mean summer A competitive sunfish, the bluegill Lepomis temperatures close to 5 degrees warmer than machrochirus, commonly occurs with pumpkinseeds native lakes. First year pumpkinseeds in their native range, but is not pumpkinseed males were observed to be present in the Iberian Peninsula. It has been sexually mature and maturation occurred earlier shown that baring the presence of this fish, morphological character release occurs in avoided any conflicts with wild salmonids and pumpkinseeds. Pumpkinseeds have been shown were stocked with a variety of species including to be trophically polymorphic having both a pumpkinseeds, , large and smallmouth “littoral” form found in the shallow water bass, and channel . Before 1999, the habitat, and a “limnetic” form that populates the funding for the program came directly from the open-water environment (Robinson et al. 1993). program as a $5.00 licensing fee, now This phenotypic plasticity accounts for 53% of combination and freshwater licensing fees the differences, where as genetics attributes provide funding. Data up until 2002 states that 14%. This plasticity allows the pumpkinseed to bluegill were still being stocked, but exploit prey in more than one habitat (Robinson pumpkinseeds stocking data was not provided et al. 1993). The limnetic morph, which feeds (Washington Department of Fish and Wildlife more predominantly on zooplankton, has smaller 2002). gaps between gillrakers, smaller paired fins, a The Department of Fish and Wildlife considers more shallow body and a longer caudal peduncle any planting of fish not approved by them as (Bhagat et al. 2006). The littoral morph’s have “unauthorized” introductions. An example given an enlarged buccal area and hypertrophied jaw is an unauthorized introduction of pumpkinseed muscles and bones in order to crush shells into a lake with a fishery in place. On (Wainwright & Richard, 1995). average, pumpkinseed fish in Washington are no longer than 6 inches (Wydoski and Whitney 1979), which diminishes their food and sport Management strategies and Control Methods value (Washington Department of Fish and Wildlife 2002). A five year old female

Pumpkinseed fish were introduced into pumpkinseed can produce anywhere from 1,950 Washington State in the early 1900’s, along with to 2,923 eggs per spawning (Wydoski and 8 other species in the sunfish family. The Whitney 1979). These abundant fry out compete purpose of the introductions was to alleviate the trout for space and food, and consume trout fishing pressure on natural fishes as well as to fry (Washington Department of Fish and provide fishing of native species for southerners, Wildlife 2002). midwesterners, and easterners who had moved Management of unauthorized fish introductions west. In 1996 the Washington State Legislature is through the use of rotenone, a natural established the Warmwater Gamefish chemical put into the lake that inhibits fish gas Enhancement Program. The goal for the exchange. In 30-60 days the lake can be program was to increase fishing opportunities restocked with the desired fish species via stocking. 80-100 lakes were chosen that (Washington Department of Fish and Wildlife Bulletin State College Forum. 2002). Roosevelt Wildlife Annual 1(1-2). 283-542. In Europe, García-Berthou and colleagues Bagenal, T.B. and Braum, E. 1971. Eggs and (2005), propose that pathways of introductions early life history. In: Methods for need to be the management focus because this assessment of fish production in fresh step has had more impact than establishment. waters, 2nd ed. (Ricker WE., ed). London: They also state that previous history of Blackwell Scientific Publications; 166-198. successful establishment is an important element Balon, E.K. 1959. Die Entwicklung des to focus on. They identify large central akklimatisiertwn Lepomis gibbosus (Linné European countries as sources and spread moves 1948) während der embryonalen Periode in initially in a southern direction, followed by a den Donauseitenwässern. Z. Fischerei 8(1- northern direction. For instance they stated that 3):1-27. most of Spain’s invaders come from southern Becker, George, C. 1983. Fishes of Wisconsin. France, therefore the next country needs to have The University of Wisconsin Press. 828- tight controls on known pathways once it’s 833. “donor” country is know to be invaded. Bhagat, Y. Fox, M.G. and Ferreira, M.T. 2006. Management of pumpkinseed in England allows Morphological differentiation in introduced government agencies to control movements and pumpkinseed Lepomis gibbosus (L.) natural dispersal of specifically pumpkinseed to occupying different habitat zones in rivers that join to the lake, but there is no law Portuguese reservoirs. Journal of Fish from moving other fish from one location to Biology. 69 (Supplement C), 79-94. another location, including species known to Breder, C.W. Rosen, D.E. 1966. Modes of carry hitchhiking pumpkinseeds (Villeneuve et reproduction in fishes. American Museum al, 2005). of Natural History. New York. 941pp Brouwer, A.B. 1925. Het invoeren van vreemde vischsoorten I. Onze Zoetwatervisscherij References 21: 17-18. Burns, J.R. 1979. The Reproductive Cycle and 2001 Sportsman Lake Stock Assessment Its Environmental Control in the Survey: The Warmwater Fish March 2003 Pumpkinseed, Lepomis gibbosus (Pisces: Community of an Isolated Island Lake 17 Centrarchidae. COPEIA (3): 449-455 Adams, C.C. and Hankinson, T.L. 1928. Carlander, K.D. 1977. Handbook of Freshwater Annotated list of Oneida Lake Fish. Biology. Vol 2, The State Press. Ames: 55-63. Chapman, W.M. 1942. Alien fishes in the waters Dembski, S., Masson,G., Monnier, D., Wagner, of the Pacific Northwest. California Fish P., and Pihan, J.C. 2006. Consequences of and Game. 28(1) :9-15 elevated temperatures on life-history traits Clark, F.W. and Keenleyside, M.H.A. 1967. of an introduced fish, pumpkinseed Reproductive isolation between the sunfish Lepomis gibbosus. Journal of Fish Biology. Lepomis gibbosus and L. macrochirus. 69, 331-346. Journal of Fish Res. Board Canada 24(3): DeWitt, T.J. 1998. Costs and limits of 495-514. phenotypic plasticity: tests with predatory- Cone, D. K., Anderson, R.C. 1977. induced morphology and life history in a Myxosporidan Parasites of Pumpkinseed freshwater snail. Journal of Evolutionary (Lepomis gibbosus L.) from Ontario. The Biology. 11, 465-480. Journal of Parasitology, Vol. 63, No. 4. pp. Elvira, B. 1995. Freshwater fishes introduced in 657-666. Spain and relationships with autochthonous Copp, G.H., Fox, M.G. and Kovac, V. 2002. species. In: Voigtlander, C.W. (Ed.), The Growth, morphology and life history traits State of the World’s Fisheries resources. of a coolwater European population of Proceedings of the World Fisheries pumpkinseed Lepomis gibbosus. Journal of Congress, Theme 3: 261-264. New Delhi: Freshwater Ecology. 4: 419-423. Oxford & IBH Publishing Co. Pvt. Copp, G.H., Fox, M.G., Przybylski, M., Emery, A. R. 1973. Preliminary comparisons of Godinho, F.N., and Vila-Gispert, A. 2004. day and night habitats of freshwater fish in Life-time growth patterns of pumpkinseed Ontario lakes. Journal of Fish Res. Board Lepomis gibbosus introduced to Europe, Canada 30(6): 761-774. relative to native American populations. García-Berthou, E. and Moreno-Amich, R. Folia Zoologica. 53,3:237-254. 2000. Food of Introduced Pumpkinseed Crossman, E.J., 1991. Introduced freshwater Sunfish: Ontogenetic Diet Shift and fishes: A review of the North American Seasonal Variation. Journal of Fish perspective with emphasis on Canada. García-Berthou, E., Alcaraz, C., Pou-Rovira, Q., Canadian Journal of. Fisheries and Aquatic Zamora, L., Coenders, G. and Feo, C. 2005. Science 48 (Suppl. 1):46-57. Introduction pathways and establishment De Groot, S.J. 1985. Introductions of non- rates of invasive aquatic species in Europe. indigenous fish species for release and Canadian Journal of Fisheries and Aquatic culture in The Netherlands. Aquaculture 46: Sciences. 62,2; Research Library pg453. 237-257. Garcia de Jalón, D. Mayo Rustarazo, M. Gallego, B. and Hervella, F. 1993. Las comununidas piscicolas de los embalses de Klaar, M., Copp, G.H., Horsfield, R. 2004. Madrid. Directrices para su gestión. Autumnal habitat use of non-native Ecología 7, 467-485. pumpkinseed Lepomis gibbosus and Godinho, F.N., Ferreira, M.T. and Cortes, R.V. associations with native fish species in 1997. The environmental basis of diet small English streams. Folia Zoologica. 53, variation in pumpkinseed sunfish, Lepomis 2; pg.189. gibbosus, and largemouth bass, Künstler, J. 1908. Amiurus nebulosus et Micropterus salmoides, along an Iberian Eupomotis gibbosus. Bulletin de la Societé river basin. Environmental Biology of d’Acclimataion. 238-244. Fishes. 50, 105-115. Lampman, B.H. 1946. The Coming of the Pond Grimaldi, E. 1972. Lago Maggiore: effects of Fishes. :177 pp. exploitation and introductions on the Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. salmonid community. Journal of Fisheries Jenkins, D.E. McAllister, and J.R. Stauffer, Research Board of Canada 29: 777- Jr. 1980. Atlas of North American 785.Biology. 57: 29-40. Freshwater Fishes. 1980(12):854 pp Gross, M.R. 1979. Cuckoldry in sunfish Looyen, A.J.L. 1948. De Nederlandsche (Lepomis Centrarchidae). Canadian Journal Heidamaatschappij en haar bemoeiingen op of Zoology. 57: 1507-1509. visserjigebied. In : De Nederlandse Gross, M.R. 1980. Sexual selection and the Heidamaatschappij 60 jaar. 152-168. evolution of reproduction strategies in Arnhem: De Nederlandsche sunfishes (Lepomis Centrarchidae) (PhD Heidamaatschappij. dissertation). Utah: University of Utah. Minchew, D.C., Sleight III,B.H. 1977. Hart Jr., C.W. and Fuller, S.J.H. 1974. Pollution Henneguya episclera sp. n. (Protozoa: ecology of freshwater invertebrates. Myxosporida), a New Myxosporidan from Academic Press, N.Y. 389pp. the Eye of the Pumpkinseed Sunfish. The Houde, E.D. and C.E. Zastrow, 1993. Journal of Parasitology, Vol. 63, No. 4. pp. Ecosystem- and taxon-specific dynamic and 667-668 energetic properties of fish larvae Mittelbach, G.G. and Chesson, P.L. 1987. assemblages. Bulliten of Marine Science. Predation risk: indirect effects on fish 53(2):290-335. populations. In Kerfoot, W.C. and Sih, A. Keith, P. and Allardi, J. 2001. Atlas des eds. Predation. University Press of New Poissons d’Eau Douce de France. Muséum England, Hanover, NH, pp315-332. National d’histoire Naturelle, Paris. 387 p. Muus, B.J. and Dahlstrøm. 1968. Guide des Roule, L. 1931. Les poisons et le monde vivant Poissons d’eau Douce et Pêche. Neuchâtel, des eaux. Tome II. Les oeufs et les nids. Switzerland: Delachaux and Niestlé. Librairie Delegrave. Paris. Nikolsky, G.V., 1961. Special Ichthyology. Simpson, J. and R. Wallace. 1978. Fishes of Jerusalem: Israel Program for Scientific Idaho. :237 pp, Translation. The Fisheries Society of the British Isles. 2006. Raveret-Wattel, C. 1900. Poissons D’Eau Life-history variation & reproductive Douce. Paris; Librairie des Sciences allocation in introduced pumpkinseed Naturelles. (Lepomis gibbosus) populations in the UK Reznick, D.A., and Endler, J.A., 1982. The 2006, Journal of Fish Biology, 69 impact of predation on life history (Supplement C), 246–265) evolution in Trinidadian guppies (Poecillia Thienemann, A. 1950. Die Binnengewässer, reticulate). Evolution 36, 160-177. Band 18 Vebreitungsgeschichte de Rios-Cardenas, O. 2003. The use of molecular Süsswassertierwelt Europas. Stugart: genetic analysis to examine mating patterns Schweizerbart. and parental care in the pumpkinseed Villeneuve, F., Copp, G.H., Fox, M.G. and sunfish (PhD dissertation). Buffalo: State Stakenas, S. 2005. Interpopulation variation University of New York at Buffalo. in growth and life-history traits of the Rios-Cardenas, O. and Webster, M.S. 2005. introduced sunfish pumpkinseed Lepomis Paternity and parental effort in the gibbosus, in southern England. Journal of pumpkinseed sunfish. Behavior Ecology. applied Ichthyoligy. 21: 275-281. July 2005: 914-921. Wainwright, P.C. and Richard, B.A., 1995. Robinson, B.W., Wilson, D.S., Margosian, A.S. Predicting patterns of prey use from and Lotito, P.T. 1993. Ecological and morphology of fishes. Environmental morphological differentiation by Biology of Fishes. 44: 97-113. pumpkinseed sunfish in lakes with out Washington Department of Fish and Wildlife. billgill sunfish. Evolutionary Ecology. 7: 2005. Warmwater Fishes of Washington. 451-464. (FM93-9) :24 Robinson, B.W., Wilson, D.S., Margosian, A.S. Welcomme, R.L. 1981 Register of international 2000. A pluralistic analysis of character transfers of inland fish species. FAO releases in pumpkinseed sunfish (Lepomis Fisheries Technical Paper 213: 120pp gibbosus). Ecology, 81(10); 2799-2812. Werner, E.E. and Hall, D.J. 1976. Niche shifts in sunfishes: experimental evidence and significance. Science 191:404-406. Wheeler, A. 1969. The fishes of the Brithis Isles and Northwest Europe. Macmillian & Co., London. Wydoski, R., S., and Whitney, R.,R. 1979. Inland Fishes of Washington. University of Washington Press. 140-141.

Current Research and Management Efforts, and Contact Information

Current research was not apparent.

Under the Washington State Aquatic Nuisance Species Watch list, pumpkinseed fish are classified as (RCW 77.60.130 2007). They are not considered a “Priority or Secondary Species of Concern”, but are on the “Regulated Species” list as the risk they pose is considered manageable relative to their benefits.. They are also on the WDFW Prohibited list (WAC 220- 12-09) as well as on the WDFW Regulated Species list (WAC 220-12-090) Outside of this there is no management plan. For more information or questions about the Warmwater Gamefish Enhancement Program, contact: Washington Department of Fish and Wildlife Attention: Angler Education Program 600 Capitol Way North, M/S 43200 Olympia WA 98501-1091