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Adaptations of Mustelinae to Winter Or, They didn’t listen to Bergmann

Adaptations of Mustelinae to Winter Or, They didn’t listen to Bergmann. Long Thin Body Types Small and medium sized What about Bergmann’s rule?. Sheryn Olson, Winter Ecology, Spring 2005. Who are the Mustelinae?. Texasferret.org. Subfamily Mustelinae. Smaller weasels & stoats

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Adaptations of Mustelinae to Winter Or, They didn’t listen to Bergmann

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  1. Adaptations of Mustelinae to WinterOr, They didn’t listen to Bergmann • Long Thin Body Types • Small and medium sized • What about Bergmann’s rule? Sheryn Olson, Winter Ecology, Spring 2005

  2. Who are the Mustelinae? Texasferret.org

  3. Subfamily Mustelinae Smaller • weasels & stoats • ferrets & polecats • mink Larger • martens • fishers

  4. Smaller Mustelinae Least weasel (Mustela nivalis) Short Tailed Weasel or European Stoat (Mustela erminea) Long Tailed Weasel (Mustela frenata)

  5. Smaller Mustelinae Black Footed Ferret (Mustela nigripes) European Polecat (Mustela putorius eversmanii) American Mink Mustela vison

  6. Larger Mustelinae • Martens • Fishers

  7. Larger Mustelinae Pine Marten Martes americana • Martens • Fishers Stone or Beech Marten Martes foines

  8. Larger Mustelinae • Martens • Fishers Fisher Martes pennanti

  9. What is Bergmann’s Rule • A smaller surface to volume ratio means a lower rate of heat loss. • The further north an endothermic species is distributed, the larger their body size. (Bergmann, 1847)

  10. Bergmann’s Rule • Is Bergmann’s rule valid? • Methods of quantification • Hypotheses to explain mechanisms • Six hypotheses (Blackburn & Hawkins, 2004)

  11. Bergmann’s Rule and Mustelinae • Is that species endothermic or homeothermic? (Rensch, 1938 and Mayr, 1956) • Is a species distributed widely? • Are comparisons inter- or intra- specific? (James, 1970 and Blackburn, et al, 1999)

  12. Bergmann’s Rule and Mustelinae • Is a species distributed widely? Mink Lariviére, 1999

  13. Bergmann’s Rule and Mustelinae • Is a species distributed widely? Long Tailed Weasel Sheffield & Thomas, 1997

  14. Bergmann’s Rule and Mustelinae • Is a species distributed widely? Short Tailed Weasel King, 1983

  15. Bergmann’s Rule and Mustelinae • Is a species distributed widely? Least Weasel M. nivalis Sheffield & King, 1994

  16. The Fight For Mayr, 1956, 1963 Rosenzweig, 1968 Paterson, 1990 Ashton, 2000, 2001 Against Rensch, 1938 McNab, 1971 Geist, 1987 Erlinge, 1987 Gould, 1997 Gaston, et. al., 1998 Meiri & Dayan, 2003

  17. Current Status • 2000, 2001: Ashton YES • 2004: Blackburn & Hawkins MAYBE • 2004: Meiri, et al MAYBE

  18. Questions for Weasels • Why don’t weasels follow Bergmann’s Rule? • What are the advantages of a small, thin body in Northern Latitudes?

  19. Physiological Adaptations • higher metabolic rate (Mustela nivalis)(Scholander, 1950) • delayed implantation of embryos M. frenata and M. erminea • pelagic color changes • fur length (Sheffield & King, 1994) M. Erminea changing pelage

  20. Behavioral Adaptations • switching daily activity patterns (Zielinski, 2000) • switching food sources (Danilov and Tumanov, 1976; Weber, 1990 Pandolfi, et al, 1996) • caching food (Weeks, 1991; Henry, et al., 1990; King, 1989; Oksanan, et al 1985) • fur lined nests

  21. Is Bergmann’s rule invalid? Morphology • Long and thin works year round • The better to prey on subnivean and burrowing mammals • The better to avoid predators by going underground • Other adaptations compensate: Behavioral and Physiological

  22. Discussion • Small Mustelinae use other strategies to maintain their energy balance in winter. • Weasel species appear not to follow Bergmann’s rule. • Or do they? Short Tailed Weasels… • (King, 1983)

  23. In Conclusion • Mustelinae illustrate it is important to consider a whole suite of adaptations. • When observing patterns, don’t confuse them with mechanisms.

  24. Selected References: The Bergmann’s Rule Controversy Ashton, Kyle G. 2001. Are ecological and evolutionary rules being dismissed prematurely? Diversity and Distributions. 7: 289-295. Ashton, K.G., Tracy, M.C. & de Queiroz, A. 2000. Is Bergmann’s rule valid for mammals? American Naturalist. 156:390 – 415. Bergmann, Carl. 1847. Ueber die Verhältnisse der Wärmeokonomie der Thiere zu ihrer Grösse. Gottinger studien. 3:595-708 Blackburn, T.M. & Gaston, K.J. and Loder, N. 1999. Geographic gradients in body size: a clarification of Bergmann’s rule. Diversity and Distributions. 5: 165-174. Blackburn, T.M. & Hawkins, B.A. 2004. Bergmann’s rule and the mammal fauna of North America. Ecography. 27: 715-724. Dayan, Tamar, Simberloff, Daniel, Tchernov, E. and Yoran YT. 1989. Inter- and intraspecific Character Displacement in Mustelids. Ecology. 70(5): 1526-1539 Erlinge, S. 1987. Why do European stoats not follow Bergmann’s Rule? Holarctic Ecology. 10: 33-39 Gaston, K.J. Blackburn, T.M. & Spicer, J. 1998. Rapoport’s rule:Time for an epitaph? Trends in Ecology and Evolution. 13: 70-74.

  25. Selected References: The Bergmann’s Rule Controversy Geist, V. 1987. Bergmann’s rule is invalid. Canadian Journal of Zoology. 65: 1035-1038. Geist, V. 1990. Bergmann’s rule is invalid: a reply to J.D. Paterson. Canadian Journal of Zoology. 68: 1613-1615. Gould, S.H. 1997. Cope’s rule as psychological artifact. Nature. 385: 199-200. Meiri S, Dayan T., Simberloff, D. 2004. Carnivores, biases and Bergmann’s rule . Biological Journal of the Linnean Society. 81 (4): 579-588. Mayr, Ernst. 1956. Geographical character gradients and climatic adaptation. Evolution. 10: 105-108. Mayr, Ernst. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Mass. Paterson, J.D. 1990. Comment – Bergmann’s rule is invalid: a reply to V. Geist. Canadian Journal of Zoology 68: 1610 – 1612. Rensch, B. 1938. Some problems of geographical variation and species-formation. Proceedings of the Linnean Society of London. 150: 275- 285 Rosenzweig, M.L. 1995. Species diversity in space and time. Cambridge University Press, Cambridge. 

  26. References Cited Ashton, Kyle G. 2001. Are ecological and evolutionary rules being dismissed prematurely? Diversity and Distributions. 7: 289-295. Ashton, K.G., Tracy, M.C. & de Queiroz, A. 2000. Is Bergmann’s rule valid for mammals? American Naturalist. 156:390 – 415. Bergmann, Carl. 1847. Ueber die Verhältnisse der Wärmeokonomie der Thiere zu ihrer Grösse. Gottinger studien. 3:595-708 Blackburn, T.M. & Gaston, K.J. and Loder, N. 1999. Geographic gradients in body size: a clarification of Bergmann’s rule. Diversity and Distributions. 5: 165-174. Blackburn, T.M. & Hawkins, B.A. 2004. Bergmann’s rule and the mammal fauna of North America. Ecography. 27: 715-724. Danilov, P.I. and I.L Tuminov. 1976. Mustelids of northwestern USSR. Institut Biologii, Karelskii Filial, Akademii Nauk SSSR, Leningrad, 256 pp. (in Russian)

  27. References Cited Dayan, Tamar, Simberloff, Daniel, Tchernov, E. and Yoran YT. 1989. Inter- and intraspecific Character Displacement in Mustelids. Ecology. 70(5): 1526-1539 Erlinge, S. 1987. Why do European stoats not follow Bergmann’s Rule? Holarctic Ecology. 10: 33-39 Geist, V. 1987. Bergmann’s rule is invalid. Canadian Journal of Zoology. 65: 1035-1038. Geist, V. 1990. Bergmann’s rule is invalid: a reply to J.D. Paterson. Canadian Journal of Zoology. 68: 1613-1615. Hall, ER. 1951. American Weasels. Univ of Kansas Publication Museum of Natural History. 4:1-466. Henry SE, Raphael MG, Ruggiero LF. 1990. Food caching and handling by marten. Great Basin Nat 50:381-383  James, F. C. 1970. Geographic size variation in birds and its relationship to climate. Ecology 51: 365-390. King, Carolyn M. 1983. Mustela erminea. Mammalian Species. 197, pp. 1-8. Lariviére, Serge.  1999.  Mustela vison. Mammalian Species. American Society of Mammalogists. No. 608. pp. 1-9. 3 fig. Distribution map of mink.

  28. References Cited Loso, H. 1999. "Mustela erminea" (On-line), Animal Diversity Web. Accessed February 24, 2005 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Mustela_erminea.html. Mayr, Ernst. 1956. Geographical character gradients and climatic adaptation. Evolution. 10: 105-108. Mayr, Ernst. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, Mass. McNab, Brian K. 1971. On the Ecological Significance of Bergmann's Rule. Ecology: Vol. 52, No. 5, pp. 845–854. Meiri S, Dayan T., Simberloff, D. 2004. Carnivores, biases and Bergmann’s rule . Biological Journal of the Linnean Society. 81 (4): 579-588. Oksanan, T, Oksanen, L, Fretwell SD. 1985. Surplus killing in the hunting strategy of small predators. Am Nat 126:328-346. Meiri S, Dayan T., Simberloff, D. 2004. Carnivores, biases and Bergmann’s rule. Biological Journal of the Linnean Society. 81 (4): 579-588.

  29. References Cited Pandolfi M, Demarinas AM & Petrov I. 1996. Fruit as a winter feeding resource in the diet of stone marten (Martes foina) in east-central Italy. Z. Säugetierk. 61 : 215-220. Paterson, J.D. 1990. Comment – Bergmann’s rule is invalid: a reply to V. Geist. Canadian Journal of Zoology 68: 1610 – 1612. Rensch, B. 1938. Some problems of geographical variation and species-formation. Proceedings of the Linnean Society of London. 150: 275- 285 Rosenzweig, M.L. 1995. Species diversity in space and time. Cambridge University Press, Cambridge. Sheffield, Steven R. and King, Carolyn M. 1994. Mustela nivalis. Mammalian Species. 454, pp. 1-10. Sheffield, SR. and Thomas, HH. 1997. Mustela frenata. Mammalian Species. 570:1-9. Scholander, PF., V Walters, WR Hock, L Irving. 1950. Body insulation of some arctic and tropical mammals and birds. Biol. Bull 99:225-236.

  30. References Cited Stevens, G. C. 1989. The latitudinal gradient in geographical range: how so many species co-exist in the tropics. Am. Nat. 133:240-256. Weeks, HP. 1991. Arboreal caching of prey by long-tailed weasel. The Prairie Naturalist. 25:39-42. Weber J. M. 1990. Seasonal exploitation of amphibians by otters (Lutra lutra ) in north-east Scotland. J. Zool. Lond. 220, 641-651. Zielinski, William J. 2000. Weasels and Martens – Carnivores in Northern Latitudes. Chapter 5, pp. 95-118 in Activity Patterns in Small Mammals. Ecological Studies. Vol. 141. S. Halle, N.C. Stenseth (Editors) http://www.inhs.uiuc.edu/dnr/fur/species/weasel.html  Global Invasive Species List: http://www.issg.org/database/species/distribution.asp?si=98&fr=1&sts=sss# (for M. erminea) http://www.washington.edu/burkemuseum/collections/mammalogy/mamwash/muer.html - distribution and maps of M. erminea Texasferret.org: http://www.texasferret.org/news/199901.shtml Family Tree University of Michigan Museum of Zoology Animal Diversity Web: http://animaldiversity.ummz.umich.edu/site/accounts/information/Mustelidae.html  Zoo page from Yugoslavia http://www.nrtonline.co.yu/zoopage/klas/mesojedi.htm– many photos.

  31. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1851: The black-footed ferret is reported and described by John James Audubon. No one will report seeing a ferret again for another 26 years.1964: A female ferret and kits are found in Mellette County in western South Dakota. They are considered perhaps the last black-footed ferrets in the world.1967: The black-footed ferret is put on the Endangered Species list.1971: Executive Order 11643 is implemented, banning the use of poisons with secondary hazards on public lands. 1972: A drowned ferret is discovered in a watering tank in Wyoming. No others are found.Nine South Dakota ferrets are captured and taken to Patuxent Wildlife Research Center in Laurel, Maryland. It is hoped that these ferrets will produce kits, but every litter is dead at birth.

  32. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1973: The Endangered Species Act is passed.1979: The last Patuxent ferret dies. The black-footed ferret is declared extinct.1981: September 25th -- A Wyoming ranch dog belonging to John and Lucille Hogg kills a black-footed ferret.October 29th -- A live black-footed ferret is spotted near Meeteetse, Wyoming. Conservationists and researchers begin an intensive search and study of wild ferrets.1984: The Meeteetse population is 129 black-footed ferrets. Plans are made to begin a captive breeding program.1985: Outbreaks of sylvatic plague and canine distemper nearly wipe out the Meeteetse ferrets.

  33. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.orgAugust 27th -- The US Fish & Wildlife Service and Wyoming Game and Fish Department decide to remove all known black-footed ferrets from their habitat in an effort to save the species. From October 1985-September 1986, 17 ferrets are taken into captivity.1986: The IUCN--World Conservation Union's Conservation Breeding Specialist Group (CBSG) conducts a workshop to develop a recovery plan for captive breeding and reintroduction of black-footed ferrets.1987: The last known ferret is captured at Meeteetse in February. These 18 captive black-footed ferrets are probably the rarest mammals on earth.A captive breeding program is initiated by the US Fish & Wildlife Service, The Wyoming Game & Fish Department, and the American Zoo & Aquarium Association. Two litters of ferret kits are born at Sybille Wildlife Research and Conservation Education Center, Wyoming to "Becky" and "Jenny". This brings the total number of black-footed ferrets in captivity to 25.

  34. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1988: In October, eight ferrets are flown to the National Zoological Park's Conservation and Research Center in Front Royal, VA to start a new breeding colony and to guard against catastrophic loss of the captive population. December, eight more ferrets are taken to the Henry Doorly Zoo in Omaha, NE."Conservation Biology of the Black-footed Ferret" (Seal et al.) is published.The "Black-footed Ferret Recovery Plan" is drafted by the US Fish & Wildlife Service.1989: 72 ferret kits are born at Sybille and 6 at Front Royal. The total ferret population is 120.1990: The Cheyenne Mountain Zoo in Colorado Springs, Colorado joins the captive breeding program.1991: Shirley Basin, in central Wyoming becomes the first black-footed ferret reintroduction site with the release of 49 juvenile ferrets.

  35. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1991: A Species Survival Plan (SSP) is developed in cooperation with the American Zoo and Aquarium Association to manage the genetic and demographic needs of the captive ferret population.The Louisville Zoological Garden in Louisville, Kentucky and the Phoenix Zoo in Phoenix, AZ join the captive breeding program.1992: Two litters of wild-born kits are reported in Shirley Basin--the first known kits born in the wild since the Meeteetse population was lost.1993: Four wild-born litters are discovered in Shirley Basin.The Toronto Zoo, Toronto, Canada joins the captive breeding program.1994: The Charles M. Russell National Wildlife Refuge in Montana, and Badlands National Park, South Dakota receive ferrets for reintroduction.An outbreak of plague spreads throughout the Shirley Basin release site and further reintroductions are postponed. A total of 228 ferrets released in Shirley Basin from 1991-1994. Small numbers of ferrets persist there today.1995: The US Fish & Wildlife Services assumes responsibility for managing the Sybille, WY breeding facility and renames it the National Black-footed Ferret Conservation Center.Ferrets are found to be highly susceptible to direct contact with plague.1996: The US Fish & Wildlife Service establishes a Black-footed Ferret Recovery Implemen-tation Team to help guide recovery efforts. The BFFRIT includes representatives from federal and state governments, Native American tribes, zoos, and conservation organiza-tions.The South Dakota ferret reintroduction effort expands onto the Buffalo Gap National Grasslands.Arizona joins the reintroduction program by testing the use of on-site "preconditioning" pens for the first time at a release site in the Aubrey Valley in northwestern Arizona.1997: Twenty-four on-site preconditioning pens are constructed by the US Forest Service on the Buffalo Gap National Grasslands. 1997 is the first year that all ferrets destined for reintroduction are preconditioned before release.A new reintroduction effort begins on the Fort Belknap Indian Reservation, Montana.1998: The captive breeding program experiences its best year ever with a record 452 ferrets born and 339 surviving to weaning. Eighteen of the weaned young are produced in on-site pens in Arizona--an important milestone.The number of ferrets available for reintroduction exceeds all previous years with ap-proximately 210 kits released to the wild in Arizona, Montana, and South Dakota.The pen breeding program expands with the addition of pen facilities built by the Turner Endangered Species Fund in New Mexico. Twenty pens are completed at a pending reintroduction site in northwest Colorado, and twenty pens are constructed at Bowdoin National Wildlife Refuge in Montana to support on-going release efforts in Montana.The US Fish & Wildlife Service plans for construction of a new breeding facility near Fort Collins, Colorado to replace the current Wyoming facility.Perhaps the most important milestone in 1998 is the fact that for the first time since the black-footed ferret recovery program began, there are more ferrets in the wild than in captivity. To date, more than 2600 ferrets have been produced in captivity.:

  36. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1995: The US Fish & Wildlife Services assumes responsibility for managing the Sybille, WY breeding facility and renames it the National Black-footed Ferret Conservation Center.Ferrets are found to be highly susceptible to direct contact with plague.1996: The US Fish & Wildlife Service establishes a Black-footed Ferret Recovery Implementation Team to help guide recovery efforts. The BFFRIT includes representatives from federal and state governments, Native American tribes, zoos, and conservation organizations.The South Dakota ferret reintroduction effort expands onto the Buffalo Gap National Grasslands.Arizona joins the reintroduction program by testing the use of on-site "preconditioning" pens for the first time at a release site in the Aubrey Valley in northwestern Arizona.

  37. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1997: Twenty-four on-site preconditioning pens are constructed by the US Forest Service on the Buffalo Gap National Grasslands. 1997 is the first year that all ferrets destined for reintroduction are preconditioned before release.A new reintroduction effort begins on the Fort Belknap Indian Reservation, Montana.1998: The captive breeding program experiences its best year ever with a record 452 ferrets born and 339 surviving to weaning. Eighteen of the weaned young are produced in on-site pens in Arizona--an important milestone.The number of ferrets available for reintroduction exceeds all previous years with ap-proximately 210 kits released to the wild in Arizona, Montana, and South Dakota.

  38. Black Footed Ferrets Black-footed Ferret Timeline ------- http://www.blackfootedferret.org1998: The pen breeding program expands with the addition of pen facilities built by the Turner Endangered Species Fund in New Mexico. Twenty pens are completed at a pending reintroduction site in northwest Colorado, and twenty pens are constructed at Bowdoin National Wildlife Refuge in Montana to support on-going release efforts in Montana.The US Fish & Wildlife Service plans for construction of a new breeding facility near Fort Collins, Colorado to replace the current Wyoming facility.Perhaps the most important milestone in 1998 is the fact that for the first time since the black-footed ferret recovery program began, there are more ferrets in the wild than in captivity. To date, more than 2600 ferrets have been produced in captivity.

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