Gasterosteus aculeatus

(Linnaeus, 1758)

Threespine Stickleback

G5Secure Found in 49 roadless areas NatureServe Explorer →
G5SecureGlobal Rank
Least concernIUCN
PSESA Status
LowThreat Impact
Identity
Unique IDELEMENT_GLOBAL.2.104745
Element CodeAFCPA03010
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryVertebrate Animal
IUCNLeast concern
Endemicoccurs (regularly, as a native taxon) in multiple nations
KingdomAnimalia
PhylumCraniata
ClassActinopterygii
OrderPerciformes
FamilyGasterosteidae
GenusGasterosteus
USESAPS
COSEWICPS:X,E,T,SC
Other Common Names
Épinoche à trois épines (FR) threespine stickleback (EN)
Concept Reference
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. American Fisheries Society, Special Publication 20. 183 pp.
Taxonomic Comments
Complex patterns of variation make taxonomic treatment difficult. This stickleback may be considered a species complex with many unique and reproductively isolated populations, subspecies or species.

Populations exist that are strictly marine, anadromous, and freshwater resident. The marine and anadromous forms have given rise to diverse resident phenotypes. Subspecies have been recognized in the past, but current scientific discussion of this species complex recognizes multiple distinct species within evolutionary radiations; current genetic research is underway to determine relationships between evolutionary groups and species before names can be assigned (Hatfield 2001a, 2001b). Lateral plate morphs of resident freshwater forms are recognized as lows (i.e., lateral plates on anterior parts of the fish only), partials (i.e., lateral plates on anterior and posterior ends of the fish with a gap between), and completes (i.e., lateral plates in a continuous row anterior to posterior). In a rare form, plates are entirely absent; G. a. williamsoni is an endangered plateless form exhibiting reduction in pelvic structure, and only occurs in drainages in southern California. Lows from the Pacific coast of North America have been called G. a. microcephalus, Pacific coast completes are G. a. aculeatus, and Penczak (1964, in Wootton 1976) designated lows from Iceland as G. a. islandicus. A plateless form occurring in Shay Creek, San Bernardino County, California, has been identified as G. a. santaeannae (or santa-annae) but is currently recognized as synonymous with G. a. williamsoni (Ross 1973, Moyle et al. 1989).

Studies of allozyme variation (Haglund et al. 1992) and mitochondrial DNA sequences (Orti et al. 1994) in Asian, North American, and European populations recognized two primary clades: (1) European, North American, and some Japanese samples, which could be divided into an (1a) Atlantic basin clade comprising the eastern North American and European populations, and a (1b) basal Pacific basin assemblage comprising western North American and some Japanese populations; and (2) a divergent group of Japanese populations. The divergent Japanese clade deserves further study and possible taxonomic recognition.

Sympatric species pairs bearing "limnetic" and "benthic" life histories and distinct morphologies have evolved in several British Columbian lake systems (Thompson et al 1997, Hatfield 2001, Hatfield and Ptolemy 2001). Some populations of these are endangered or already extinct (Wood 2003).

Several low-lying lakes and streams in the Cook Inlet area contain rare and evolutionarily divergent populations of G. aculeatus including three populations polymorphic for lateral plate morphs, several populations polymorphic for pelvic armor morphs, one lake containing 2 freshwater morphs of the species (a benthic and a limnetic feeder), and one lake containing both anadromous and resident freshwater forms of the species (von Hippel, pers. comm.). Bell and Orti (1994) viewed divergent populations in freshwater habitats around Cook Inlet as parts of an endemic radiation warranting special consideration for conservation as a unit.
Conservation Status
Rank MethodExpertise without calculation
Review Date2017-12-27
Change Date1996-09-20
Edition Date2008-01-04
Edition AuthorsHammerson, G.
Threat ImpactLow
Range Extent>2,500,000 square km (greater than 1,000,000 square miles)
Number of Occurrences> 300
Rank Reasons
Widespread, abundant, secure.
Range Extent Comments
Range encompasses coastal waters of Eurasia, Iceland, Greenland, eastern Asia, and North America. In North America, this fish ranges from Alaska to Baja California on the west coast, from Baffin Island and the west side of Hudson Bay to Chesapeake Bay, Virginia, along east coast, and it occurs also in inland areas (including Lake Ontario) along both coasts. Sometimes it occurs in the open ocean. The species has been introduced and is established in certain areas of California, Massachusetts, and the Great Lakes (lakes Huron, Michigan, Erie, and Superior) (Fuller et al. 1999, Stephenson and Momot 2000). It also has been introduced in Europe, Iceland, Greenland, and the Pacific coast of Asia (Page and Burr 2011).
Occurrences Comments
This species is represented by a very large number of occurrences (subpopulations).
Threat Impact Comments
On a range-wide scale, no major threats are known.

Locally, threats include human impacts on spawning habitat and water quality, stocking or other introduction of non-native, predatory fishes, including introductions of certain forms of G. aculeatus itself.

Craig (1984) noted that large-scale industrial and petroleum extraction development in the Beaufort Sea could cause direct mortality as a result of intake of juvenile fishes with seawater and indirectly harm populations by altering coastal habitat, including water circulation patterns.

The California Department of Fish and Game (CDFG) recognized that increasing development along and recreational uses of the Santa Clara River were threats to the existence of the endangered G. a. williamsoni population (CDFG 1974).

In British Columbia, beaver activity has resulted in fluctuating water levels that have reduced access to some spawning sites used by the giant stickleback (currently recognized as G. aculeatus, but considered by some scientists to be a separate species, this fish is black in color and more than twice as long as threespine stickleback; occurs only in Mayer Lake, Queen Charlotte Islands; see sources in Rubidge 2000, Species at Risk Canada 2004). Increasing human recreational use of lake habitat also threatens this population (Species at Risk Canada 2004).

In Alaska and elsewhere, non-native species such as the northern pike (Esox lucius) and stocked salmonids may threaten sticklebacks through predation and competition for juvenile food resources (Hatfield 2001a, Hatfield and Ptolemy 2001, Foster et al. 2003, Wood 2003).

Hybridization between different forms of G. aculeatus threatens the unique genetic characteristics of specific populations recognized as rare or divergent; extensive hybridization between the native G. a. williamsoni (unarmored) and introduced G. a. microcephalus (armored) forms in California contributed to declines in the now endangered G. a. williamsoni (CDFG 1974, Moyle 1976b, Fuller 2005).
Ecology & Habitat

Description

A small (to 10 cm), laterally compressed, scaleless fish with a very narrow caudal peduncle, 3 (rarely 2 or 4) dorsal spines (the last very short) followed by a soft dorsal fin with 14-16 rays, and large eyes; pelvic fin small, with 1 spine (1 cusp at base) and 1 ray; bony keel along each side of caudal peduncle; 0-30 bony plates on each side (freshwater populations have fewer plates than do marine populations); silver-green to brown above, sides silver, often with dark mottling; large males are black and often have red on fins; breeding males have blue sides, bright red belly and lower sides, and bright blue or green eyes; large females have a pink throat and belly; colors often are subdued in inland populations (Page and Burr 1991, Moyle 1976).

Bourgeois et al. (1994) found that skeletal reduction in Alaska was related to ion composition of lake water, presence of other fishes, and local gene flow.

Diagnostic Characteristics

Differs from other sticklebacks in having fewer dorsal spines (usually 3 vs. 4 or more).

Habitat

In freshwater areas, this fish typically inhabits quiet weedy pools and backwaters, or occurs among emergent plants at stream edges, over bottoms of sand and mud (Lee et al. 1980, Page and Burr 2011). Marine populations apparently are pelagic, usually staying close to shore. In some lakes, two morphologically and ecologically distinct forms may occur, differing in habitat (one littoral, the other mainly limnetic). Eggs are deposited in freshwater in a nest of plant material made by the male on the bottom in shallow water.

Ecology

Forms loose schools except when spawning (Moyle 1976).

Available density estimates include 7-28 fish/m² in suitable habitat in Wales, 24-63 fish/m² in northwestern England, 2 fish/m² in Kamchatka, Russia, and 4-21 fish/m² in British Columbia (see sources in Wootton and Smith 2000).

Reproduction

Spawns in spring and summer. In most populations, most adults are 1-2 years old, do not live beyond 4 years, and presumably die at the end of their first breeding season. Reimchen (1992) described a population in Drizzle Lake, Queen Charlotte Islands, British Columbia, in which individuals lived up to 8 years. Male guards eggs and fry. Eggs hatch in about a week. Nest may contain eggs of several females.
Other Nations (2)
United StatesN5
ProvinceRankNative
MichiganSNANo
New YorkS4Yes
Rhode IslandS4Yes
OregonS4Yes
New HampshireS5Yes
MassachusettsS4Yes
ConnecticutSNRYes
PennsylvaniaS1Yes
WisconsinSNANo
AlaskaS5Yes
New JerseyS4Yes
MaineS5Yes
VirginiaS4Yes
MarylandSNRYes
WashingtonS5Yes
IndianaSNANo
DelawareS4Yes
CaliforniaSNRYes
CanadaN5B,N5N
ProvinceRankNative
QuebecS5Yes
AlbertaSNANo
New BrunswickS5Yes
Yukon TerritorySNANo
Northwest TerritoriesS4Yes
LabradorS5B,SNRN,SNRMYes
ManitobaS2Yes
OntarioS4Yes
NunavutSUYes
Prince Edward IslandS5Yes
Island of NewfoundlandS5B,SNRN,SNRMYes
British ColumbiaS5Yes
Nova ScotiaS5Yes
Threat Assessments
ThreatScopeSeverityTiming
No known threats

Roadless Areas (49)
Alaska (31)
AreaForestAcres
Bering LakeChugach National Forest965,076
Boston BarChugach National Forest53,617
ChichagofTongass National Forest555,858
Chilkat-West Lynn CanalTongass National Forest199,772
Chugach-12Chugach National Forest8,116
Chugach-13Chugach National Forest13,337
College FiordChugach National Forest1,130,818
Copper River WetlandsChugach National Forest85,972
Douglas IslandTongass National Forest28,065
Fidalgo-GravinaChugach National Forest257,968
Freshwater BayTongass National Forest44,933
Game CreekTongass National Forest54,469
GravinaTongass National Forest37,381
Johnson PassChugach National Forest152,508
Juneau UrbanTongass National Forest101,581
Juneau-Skagway IcefieldTongass National Forest1,187,268
KartaTongass National Forest52,117
Kenai LakeChugach National Forest213,172
Kenai MountainsChugach National Forest306,600
KogishTongass National Forest65,216
MadanTongass National Forest68,553
Nellie JuanChugach National Forest713,697
North KruzofTongass National Forest33,146
North RevillaTongass National Forest215,430
ResurrectionChugach National Forest224,615
RevillaTongass National Forest29,298
Roaded DonutChugach National Forest968
Sheridan GlacierChugach National Forest224,683
Taku-SnettishamTongass National Forest664,928
Tenakee RidgeTongass National Forest20,527
Windham-Port HoughtonTongass National Forest161,952
California (11)
AreaForestAcres
Dry LakesLos Padres National Forest17,043
Glass MountainInyo National Forest52,867
Magic MountainAngeles National Forest15,542
MatilijaLos Padres National Forest5,218
NordhoffLos Padres National Forest12,031
Orleans Mtn. BSix Rivers National Forest17,183
Red MountainAngeles National Forest8,034
San JoaquinSierra National Forest22,474
Sawmill - BadlandsLos Padres National Forest51,362
Sespe - FrazierLos Padres National Forest106,910
White LedgeLos Padres National Forest18,632
Oregon (1)
AreaForestAcres
TahkenitchSiuslaw National Forest5,799
Washington (6)
AreaForestAcres
Higgins MountainMt Baker-Snoqualmie National Forest13,185
Jupiter RidgeOlympic National Forest10,148
Mt. Baker WestMt Baker-Snoqualmie National Forest25,390
South QuinaultOlympic National Forest11,081
TaneumWenatchee National Forest26,140
Thorp Mtn.Wenatchee National Forest22,717
References (62)
  1. Bakker, T. C. M., and P. Sevenster. 1988. Plate morphs of <i>Gasterosteus aculeatus</i> Linnaeus (Pisces: Gasterosteidae): comments on terminology. Copeia 1988:659-663.
  2. Bell, M. A., and G. Orti. 1994. Pelvic reduction in threespine stickleback from Cook Inlet Lakes: geographical distribution and intrapopulation variation. Copeia 1994:314-325.
  3. Bell, M. A., and S. A. Foster. 1994. The evolutionary biology of the threespine stickleback. Oxford Univ. Press, New York. xii + 571 pp.
  4. Bell, M.A., G. Orti, J.A. Walker and J.P. Koenings. 1993. Evolution of pelvic reduction in threespine stickleback fish: a test of competing hypotheses. Evolution 47:906-914.
  5. Bourgeois, J.F., D.M. Blouw, J.P. Koenings and M.A. Bell. 1994. Multivariate analysis of geographic covariance between phenotypes and environments in the threespine stickleback, <i>Gasterosteus aculeatus</i>, from the Cook Inlet area, Alaska. Canadian Journal of Zoology 72:1497-1509.
  6. Brown, Larry (U.S. Geological Survey, WRD). 2000. Review and annotation of fish watershed distribution maps. Review requested by Anthony E. Zammit, TNC. June 2000.
  7. California Department of Fish and Game (CDFG). 1974. At the crossroads 1974: a report on California's endangered and rare fish and wildlife. Department of Fish and Game, Sacramento, CA.
  8. Coad, B. W. 1981. A bibliography of the sticklebacks. Syllogeus 35:1-142.
  9. Cooper, E. L. 1983. Fishes of Pennsylvania and the northeastern United States. Pennsylvania State University Press, University Park. 243 pp.
  10. Craig, P.C. 1984. Fish use of coastal waters of the Alaskan Beaufort Sea: a review. Transactions of the American Fisheries Society 113:265-282.
  11. Daniels, Robert (Biological Survey, New York State Museum). 2000. Review and annotation of fish watershed distribution maps. Review requested by Anthony E. Zammit, ABI. June 2000.
  12. Deagle, B. E., T. E. Reimchen, and D. B. Levin. 1996. Origins of endemic stickleback from the Queen Charlotte Islands: mitochondrial and morphological evidence. Canadian Journal of Zoology 74:1045-1056.
  13. Foster, S.A., J.A. Baker and M.A. Bell. 2003. The case for conserving threespine stickleback populations: protecting an adaptive radiation. Fisheries 28:10-18.
  14. Fuller, P. 2005. <i>Gasterosteus aculeatus</i>. USGS Nonindigenous Aquatic Species Database, Gainesville, FL. Available online at: http://nas.er.usgs.gov/queries/FactSheet.asp?SpeciesID=702. Accessed 6/9/05.
  15. Fuller, P. L., L. G. Nico, and J. D. Williams. 1999. Nonindigenous fishes introduced into inland waters of the United States. American Fisheries Society, Special Publication 27. x + 613 pp.
  16. Greenbank, J., and P.R. Nelson. 1959. Life history of the threespine stickleback <i>Gasterosteus aculeatus </i>Linnaeus in Karluk Lake and Bare Lake, Kodiak Island, Alaska. Fishery Bulletin 153, Fishery Bulletin of the Fish and Wildlife Service 59:537-559.
  17. Haglund, T. R., D. G. Buth, and R. Lawson. 1992. Allozyme variation and phylogenetic relationships of Asian, North American, and European populations of the ninespine stickleback, <i>Pungitius pungitius</i>. Pages 438-452 in R.L. Mayden, editor. Systematics, historical ecology, and North American freshwater fishes. Stanford University Press, Stanford, California. xxvi + 969 pp.
  18. Halliwell, David B. (Maine Department of Environmental Protection). 2000. Review and annotation of fish watershed distribution maps. Review requested by Anthony E. Zammit, ABI. June 2000.
  19. Hatfield, T. 2001a. Status of the stickleback species pair, <i>Gasterosteus </i>spp., in Hadley Lake, Lasqueti Island, British Columbia. Canadian Field-Naturalist 115(4):579-583.
  20. Hatfield, T. 2001b. Status of the stickleback species pair, <i>Gasterosteus </i>spp., in the Vananda Creek watershed of Texada Island, British Columbia. Canadian Field-Naturalist 115(4):584-590.
  21. Hatfield, T., and J. Ptolemy. 2001b. Status of the stickleback species pair, <i>Gasterosteus</i> spp. in Paxton Lake, Texada Island, British Columbia. Can. Field-Nat. 115(4):591-596.
  22. Hynes, H. B. N. 1950. The food of the fresh-water sticklebacks, <i>Gasterosteus aculeatus</i> and <i>Pygosteus pungitius</i>, with a review of methods used in studies of the food of fishes. Journal of Animal Ecology 19(1):182-193.
  23. Jenkins, R. E., and N. M. Burkhead. 1994. Freshwater fishes of Virginia. American Fisheries Society, Bethesda, Maryland. xxiii + 1079 pp.
  24. Krueger, William (Department of Biological Sciences, University of Rhode Island). 2000. Review and annotation of fish watershed distribution maps. Review requested by Anthony E. Zammit, ABI. February and March 2000.
  25. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina State Museum of Natural History, Raleigh, North Carolina. i-x + 854 pp.
  26. Master, L. L. 1996. Synoptic national assessment of comparative risks to biological diversity and landscape types: species distributions. Summary Progress Report submitted to Environmental Protection Agency. The Nature Conservancy, Arlington, Virginia. 60 pp.
  27. Master, L. L. and A. L. Stock. 1998. Synoptic national assessment of comparative risks to biological diversity and landscape types: species distributions. Summary Report submitted to Environmental Protection Agency. The Nature Conservancy, Arlington, VA. 36 pp.
  28. McLennan, D. A. 1993. Phylogenetic relationships in the Gasterostidae: an updated tree based on behavioral characters with a discussion of homoplasy. Copeia 1993:318-326.
  29. McPhail, J. D. 1984. Ecology and evolution of sympatric sticklebacks (<i>Gasterosteus</i>): morphological and genetic evidence for a species pair in Enos Lake, British Columbia. Canadian Journal of Zoology 62:1402-1408.
  30. McPhail, J. D. 1989. Status of the Enos Lake stickleback species pair, <i>Gasterosteus</i> spp. Canadian Field-Naturalist 103:216-219.
  31. McPhail, J. D. 1992. Ecology and evolution of sympatric sticklebacks (<i>Gasterosteus</i>): evidence for a species-pair in Paxton Lake, Texada, Island, British Columbia. Canadian Journal of Zoology 70:361-369.
  32. Metcalfe, C. D. 1989. Tests for predicting carcinogenicity in fish. Reviews in Aquatic Sciences 1(1):111-129.
  33. Moyle, P. B. 1976a. Inland fishes of California. University of California Press, Berkeley, California. 405 pp.
  34. Moyle, P. B. 1976b. Fish introductions in California: history and impact on native fishes. Biological Conservation 9:101-118.
  35. Moyle, P. B. 2002. Inland fishes of California. Revised and expanded. University of California Press, Berkeley. xv + 502 pp.
  36. Moyle, P. B., J. E. Williams, and E. D. Wikramanayake. 1989. Fish species of special concern of California. Final report submitted to California Dept. of Fish and Game, Inland Fisheries Division, Rancho Cordova. 222 pp.
  37. Nelson, J. S., E. J. Crossman, H. Espinosa-Perez, L. T. Findley, C. R. Gilbert, R. N. Lea, and J. D. Williams. 2004. Common and scientific names of fishes from the United States, Canada, and Mexico. American Fisheries Society, Special Publication 29, Bethesda, Maryland. 386 pp.
  38. O'Reilly, P., T.E. Reimchen, R. Beech and C. Strobeck. 1993. Mitochondrial DNA in <i>Gasterosteus</i> and Pleistocene glacial refugium on the Queen Charlotte Islands, British Columbia. Evolution 47(2):678-684.
  39. Orti, G., M.A. Bell, T.E. Reimchen, and A. Meyer. 1994. Global survey of mitochondrial DNA sequences in the threespine stickleback: evidence for recent migrations. Evolution 48:608-622.
  40. Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes: North America north of Mexico. Houghton Mifflin Company, Boston, Massachusetts. 432 pp.
  41. Page, L. M., and B. M. Burr. 2011. Peterson field guide to freshwater fishes of North America north of Mexico. Second edition. Houghton Mifflin Harcourt, Boston. xix + 663 pp.
  42. Page, L. M., H. Espinosa-Pérez, L. T. Findley, C. R. Gilbert, R. N. Lea, N. E. Mandrak, R. L. Mayden, and J. S. Nelson. 2013. Common and scientific names of fishes from the United States, Canada, and Mexico. Seventh edition. American Fisheries Society, Special Publication 34, Bethesda, Maryland.
  43. Page, L. M., K. E. Bemis, T. E. Dowling, H.S. Espinosa-Pérez, L.T. Findley, C. R. Gilbert, K. E. Hartel, R. N. Lea, N. E. Mandrak, M. A. Neigbors, J. J. Schmitter-Soto, and H. J. Walker, Jr. 2023. Common and scientific names of fishes from the United States, Canada, and Mexico. Eighth edition. American Fisheries Society (AFS), Special Publication 37, Bethesda, Maryland, 439 pp.
  44. Reimchen, T.E. 1984. Status of unarmoured and spine-deficient populations (Charlotte unarmoured stickleback) of threespine stickleback, <i>Gasterosteus</i> sp., on the Queen Charlotte Islands, British Columbia. Canadian Field-Naturalist 98(1):120-126.
  45. Reimchen, T. E. 1992. Extended longevity in a large-bodied stickleback, <i>Gasterosteus</i>, population. Canadian Field-Naturalist 106(1):122-125.
  46. 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. American Fisheries Society, Special Publication 20. 183 pp.
  47. Ross, S.T. 1973. The systematics of <i>Gasterosteus aculeatus </i>(Pisces: Gasterosteidae) in central and southern California. Contributions in Science 243. Los Angeles Natural History Museum, Los Angeles, CA.
  48. Rubidge, E. 2000. Variability within the <i>Gasterosteus </i>species complex in coastal British Columbia. Royal British Columbia Museum, Victoria, B.C.
  49. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada, Bulletin 184. 966 pp.
  50. Smith, C. L. 1985. The inland fishes of New York State. New York State Department of Environmental Conservation. Albany, New York, xi + 522 pp.
  51. Species at Risk Canada. 2004. Species at risk - Giant Stickleback. Available online at: http://www.speciesatrisk.gc.ca/search/speciesDetails_e.cfm?SpeciesID=97. Accessed 6/13/05.
  52. State Natural Heritage Data Centers. 1996a. Aggregated element occurrence data from all U.S. state natural heritage programs, including the Tennessee Valley Authority, Navajo Nation and the District of Columbia. Science Division, The Nature Conservancy.
  53. Stephenson, S. A. and W. T. Momot. 2000. Threespine, <i>Gasterosteus aculeatus</i>, and fourspine, <i>Apeltes quadracus</i>, sticklebacks in the Lake Superior basin. Canadian Field-Naturalist 114(2):211-216.
  54. Thompson, C. E., E. B. Taylor, and J. D. McPhail. 1997. Parallel evolution of lake-stream pairs of threespine stcklebacks (<i>Gasterosteus aculeatus</i>) inferred from mitochondrial DNA variation. Evolution 51:1955-1965.
  55. U.S. Fish and Wildlife Service (USFWS). 2002. Endangered and Threatened Wildlife and Plants; designation of critical habitat for the Unarmored Threespine Stickleback. Federal Register 67(180): 58580-58582 (17 September 2002).
  56. U.S. Fish and Wildlife Service (USFWS). 2005. Species profile for unarmored threespine stickleback. U.S. Fish and Wildlife Service Endangered Species. Available online at: http://ecos.fws.gov/species_profile/SpeciesProfile?spcode=E00X. Accessed 6/9/05.
  57. von Hippel, F. 2005. Personal communication via email between Frank von Hippel, University of Alaska Anchorage, and Jodi McClory, Alaska Natural Heritage Program regarding divergent threespine stickleback populations occurring in the Cook Inlet area, AK. July 28, 2005.
  58. Whittier, Thomas R. 2000. Review and annotation of fish watershed distribution maps. Review requested by Anthony E. Zammit, ABI. March 2000.
  59. Wood, P. M. 2003. Will Canadian policies protect British Columbia's endangered pairs of sympatric sticklebacks? Fisheries 28(5):19-26.
  60. Wootton, R. J. 1976. The biology of sticklebacks. Academic Press, London. 387 pp.
  61. Wootton, R. J. 1984. A functional biology of sticklebacks. University of California Press, Berkeley, California. 265 pp.
  62. Wootton, R.J., and C. Smith. 2000. A long-term study of a short-lived fish: the demography of <i>Gasterosteus aculeatus</i>. Behaviour 137:981-997.