1. Using Historical Information in Wildlife Science: A Personal Journey William B. Krohn, Professor Emeritus Department of Wildlife, Fisheries, and Conservation Biology

2. On History and Wildlife Science “Research programs pay too little attention to the history of wildlife … We do not yet appreciate how much historical evidence can be dug up, or how important it can be in the appraisal of contemporary ecology.” ─ Aldo Leopold (1933)

3. Presentation Outline Philosophy of Science Promises and Pitfalls Published Studies Potential Studies Take Home Message

4. Hypothesis Testing A scientific hypothesis is a testable, fact-based explanation of a natural phenomenon; it is not an assumption. In science, a hypothesis continues to be viewed as a plausible until rejected. Under the scientific method what is not rejected (i.e., “accepted”) often changes as new information is accumulated through time. Science is a process – a quest for truth.

5. Some Sources of Historical Wildlife Information Fur company ledgers and shipping records (e.g., Hudson Bay Co., British Colonial Office) Writings of the early European explorers and merchants Journal and diaries Sporting periodicals, newspapers, and popular books book Civil and natural histories Scientific papers and books

6. Pitfalls Observers are often unknown to today’s scientists and thus easy to dismiss as ‘unreliable.’ Observations are almost always descriptive and incidental, not done via an experimental design, and thus easily dismissed as ‘anecdotes’ and ‘biased.’ Information can be difficult to locate, and adequately assessing historical information can require specialized knowledge (sometimes calling for the help of historians and other specialists).

7. Know Your Observers 2005 2010

8. Promises Establish reference conditions for wildlife communities against which management and/or restoration goals can be judged. Document the composition of wildlife communities in the past to assess the degree of departures in today’s communities (descriptive). To understand the process(es) that led to the extirpation, decline, or increase of a wildlife species (predictive).

9. Example 1 – Canada Lynx Listing Under the Federal Endangered Species Act (ESA) in the Late-1990s Should Canada Lynx in Maine be listed under the Federal ESA? Did lynx occur historically in Maine? If yes, where these resident, breeding animals or non-breeding subadults dispersing into Maine from Quebec?

10. Historical Distribution of Canada Lynx in Eastern North America (from Seton 1929)

11. Data Sources 18th & 19 th century fur-trade records 19 th century state scientific surveys 19 th & early 20 th century scientific/technical literature 19 th & early 20 th century sporting periodicals and books, and personal diaries

12. Nomenclature Terms used for Lynx canadensis –Canada lynx, bay lynx, northern lynx, gray lynx, loup cervier, lucivee, lusern, luceret, bobcat, wildcat, ounce, catamount, Indian devil, lunxus, wolverene Terms used for L. rufus –Bay lynx, bobcat, wildcat, common wildcat, lynx, ounce, wolverene from Thompson (1853: 36 & 37)

13. New York’s Scientific Survey

14. “Lynx were so abundant that several hundred skins were sold in this market every year till about the last of the war [i.e. 1864-1865], when in a short time all had left, so that not a single skin was offered for several years. Then they returned in such numbers that within a few years after they first came, I was buying some 200 yearly.” Manly Hardy (1907) on Canada Lynx in Maine

15. Canada Lynx in Western Maine a “When I first moved to the Mollychunkemunk Lake [Upper Richardson Lake in the Rangeley Lakes region, in the early 1840s] I trapped and killed forty-nine of these animals in one hunting season, and since that time have killed a great many each year, but have not kept account of the number…” ____________________________________________ a - J. G. Rich. 1862. “Notes upon certain mammals in Maine.” pp. 134-140 in E. Holmes, ed. The Second Annual Report on the Natural History and Geology of the State of Maine. Augusta, Maine.

16. Cananda Lynx Shull from Umbagog Lake now at Harvard University’s MCZ

17. Summary of Maine Results 118 records of 509 lynx, 1833-1999 –104 records of 181 lynx township or finer –15 records of 187 lynx multi-township areas –43 records of 141 lynx statewide Records of 39 kittens representing 21 litters 1864-1999

18. Maine Lynx Observations, 1833-1999

19. Maine to New York In the early 1800s, a continuous distribution through- out the interior forests. The distribution became more discontinuous in the late 1800s and by the 1900s was limited to the highest elevations (e.g., White Mountains in New Hampshire) and western through northern Maine Today, lynx are absent as a breeding species in the White Mountains and in Maine are confined to the northwestern corner of the state. Hoving, C. L., R. A. Joseph, and W. B. Krohn. 2003. Recent and historical distributions of Canada lynx in Maine and the Northeast. Northeastern Naturalist 10(4): 363-382.

20. Southern Extent Northcentral Pennsylvania – about 20 records for Canada lynx from the early 1800s. (S. M. Rhodes a. 1903. Mammals of Pennsylvania and New Jersey: A Biographic, Historic, and Descriptive Account). ________________________________ a - Rhodes was associated with the Philadelphia Academy of Science, and a leading field biologist of his day in this region.

21. Example 1 - Conclusions In the 1800s and early 1900s, Canada Lynx were relatively common in Maine (for a carnivore), and populations waxed and waned. During this period, lynx were most common on the Moosehead Plateau of western and northern Maine (rarer near the coast.) Maine historically, there was a viable breeding population of lynx in Maine. Previous to 1900 lynx were distributed as far south as central Pennsylvania, but today the species’ eastern U.S. range is centered in northwestern Maine.

22. For Additional Details Hoving, C. L., R. A. Joseph, and W. B. Krohn. 2003. Recent and historical distributions of Canada lynx in Maine and the Northeast. Northeastern Naturalist 10 (4): 363-382. Online at: http://works.bepress.com/william_krohn/

23. Occurrences of Canada Lynx, 1987-1999 (n = 1,150) (from Hoving et al. 2005)

24. Best Model: (CC = 94%; n = 278) Snowfall (+) Deciduous (-) The Present – Potential Regional Occurrence of Canada Lynx (Hoving et al. 2005. Journal of Wildlife Management, 69: 747)

25. The “Good Old” Days

26. Example 2 – Forest Weasels (illustrations from Newhouse, 1893 & 1895) Fisher American marten

27. TRADITIONAL EXPANATIONS FOR RANGE CONTRACTIONS “Most suitable habitat [for the marten] has been lost throughout the southern primordial range as a result of land clearance. … In some areas where adequate habitat persisted, overtrapping caused local extirpation.” ─ Strickland and Douglas (1987: 532) “American martens and fishers reached their nadir [distributional low-point] early in this century owning to overexploitation for fur and habitat loss.” ─ Powell (1994: 11)

28. Fisher, Marten, and Snow Inter-Relationships Hypotheses published by Krohn et al. (1995) proposed that deep snow limits the northern distribution of fishers, whereas high fisher populations (via predation) limits the southern distribution of marten. If true, then historical distributions of these two species – given climate warming after the Little Holocene – should have shifted over eastern North America in a predictable fashion.

29. Recent Research Related to Snow/Fisher and Fisher/Marten Hypotheses Deep snow limits fisher – eastern marten populations correlated with deep snow – Carroll (2007) *high quality fisher habitat has low snowfall – Carr et al. (2007) northern Idaho – Albrecht et al. (2009) High fisher populations limit marten – Palomares and Caro (1999) *northcentral Maine – Hodgman et al. (1997), Payer (1999) northern Wisconsin – McCann et al. (2009)

30. STUDY OBJECTIVES (1)Determine whether or not the spatial and temporal patterns in the historic records are consistent with predictions from hypotheses proposed to explain the contemporary distributions of fishers and martens, and (2) Assess the two traditional explanations of changes in the historical distributions of fishers and martens in eastern North America.

31. STUDY APPROACH Given that the climate was colder in the 1700s and 1800s in eastern North America than today, and deep snowfall occurred further south, then use (a) the snow/fisher and fisher/marten hypotheses to predict general patterns of Martes distributions, and (b) historic occurrence records to evaluate these predictions.

32. THE LITTLE ICE AGE (definition from Mayewski and White 2002) A cool period lasting from approximately 1400 to the early 1900s. This was also a prolonged period of increased storms, especially in the higher latitudes.

33. THE LITTLE ICE AGE (from Imbrie and Imbrie 1979: 181)

34. STUDY PREDICTIONS (1) Both fishers and martens would be farther south than today; and like today, fishers would occur south of martens; (2) The distribution of martens in New England and eastern Canada would have been more continuous than today; (3) Areas of high marten populations would have fewer or no fishers (and vice versa); and (4) As climate moderated and snowfalls declined, marten would retreat north and fisher populations would increase in the newly created low-snowfall forestlands. [not discussed today due to time but limitations].

35. PREDICTIONS 1 Historical Southern Limits (from Seton 1909) Fisher Marten

36. PREDICTION 2 – Discontinuous vs. Continuous Distributions “The early records show that both the fisher (Mustela Pennanti) and the marten (Mustela Americana) were common inhabitants of not only the whole of New England, but also of the Atlantic States generally, as far southward as Virginia (excepting possibly a narrow belt along the seaboard), and even farther southward along the Alleghenies.” J. A. Allen. 1876. “The Former Range of Some New England Carnivorous Mammals,” The American Naturalist, 10: 708-715.

37. PREDICTION 3 – Inverse Relative Abundance of the Two Species Approach: Use ratio of martens to fishers in the fur harvest to make comparisons in (a) time, and (b) space.

38. Truck House Record from Machias, Maine – June 1777

39. Pelts Exported from Virginia, 1699-1715 (data from the British Colonial Office) Year Fisher Marten Wolverine 1699 163 5 7 1700 106 0 0 1701 70 0 0 1702 463 0 0 1703 5 0 0 1704 58 58 4 1705 108 112 0 1706 97 4 0 1707 0 0 0 1708 416 0 0 1709 496 12 0 1710 18 35 0 1711 1,100 76 0 1712 151 1,130 0 1713 0 0 0 1714 90 0 0 1715 14 0 0 Mean 197 179 1 Total 3,355 1,432 11

40. Ratios of Marten to Fisher in Historic Harvest Data Southern Northern (Virginia, 1699-1715) (Maine & New Brunswick, 1764-1778) Predicted Low High Actual 2.3 24.8 (n) (4,787) (6,821)

41. Marten to Fisher Ratios in Maine and New Brunswick, 1764-2001 Period Maine New Brunswick (source) (n) (n) 1764-1778 24.8 a (truck houses) (6,821) 1850-early 1900s 5.5 11.1 (trapping records) (260) (193) 1997-2001 1.6 3.6 (harvest records) (32,000) (16,452) _____________________________________ a – Maine sample (n = 373) from near the ME/NB border, so it was pooled with the sample from New Brunswick (n = 6,448).

42. Example 2 - Conclusions Both species occurred south of current ranges, with fishers occurring south of martens. ACCEPTED Martens were continuously distributed than currently. ACCEPTED Areas supporting high marten populations had low fisher populations (and vice versa). ACCEPTED

43. For Additional Details Krohn, W. B. 2012. Distribution changes of American martens and fishers in eastern North America, 1699–2001. Chapter 4, pages 58-73 in Biology and Conservation of Martens, Sables, and Fishers: A New Synthesis. Cornell University Press, Ithaca, New York. Online at: http://works.bepress.com/william_krohn/

44. Potential Historical Wildlife Studies 1) The ever-shifting wildlife populations of eastern North America. 2) How many wolf species historically inhabited the Northeast? 3) Did the European red fox replace the North American red fox in eastern North America?

45. Major Color Phases of Wild Red Foxes (from Obbard 1987) Left to Right: cross, red, silver

46. Potential Genetic Comparisons Eastern North American Specimens – earliest museum vs. contemporary specimens. Earliest museum specimens – Northeastern U.S.A. vs. Southeastern U.S.A. Contemporary English specimens vs. contemporary North American specimens. Contemporary North American specimens north vs. south of the St. Lawrence River.

47. On History and Wildlife Science “Far from being an esoteric concern, the development of an historic sensibility ought to be considered fundamental to conservation biology [and wildlife management]. In fact, any teaching or practice of conservation biology that neglects history is incomplete.” ─ C. Meine. 1999. It’s about time: conservation biology and history. Conservation Biology, 13: 1-13.

48. Acknowledgments Colleagues in the University of Maine’s Wildlife Program, especially Drs. Daniel J. Harrison and Aram J. K. Calhoun, and Christopher L. Hoving. Pauleena M. MacDougall, Maine Folklife Center. Michael Alpert, University of Maine Press. Ellen Conant Krohn, my life-time supporter and partner. Dr. Ralph S. Palmer (deceased), Zoologist, New York State Museum.

49. Comments or Questions ?

51. Major Color Phases of Red Foxes in the Wild (from Voigt 1987) Major Color Phase NA Occurrence red 45-75 % silver 2-17% cross variable _______________________________________________________ Silver and cross “rarely occur south of Canada.” The silver color phase occurs only in North America. Throughout the Old World, almost all red foxes are the red color phase.

52. Eastern North America’s Ever Changing Wildlife Populations Moose White-tailed deer Woodland caribou Gray wolf Red wolf Canada lynx Bobcat American marten Fisher Wolverine Red fox Gray fox __________________________________________________________________________________________________________________________________________ Geographic distributions of all species on the left positively correlated with deep snowfall.

53. Historical Evidence of Two Species of Wolves in Maine 1)Josselyn and other 17 th century observers mention two types of wolves (e.g., see Record 11 in EMW). 2) Many written records from the mid- to late-1800s of wolves coming back but too small to kill cows or moose, but they readily took sheep and deer) (see Record 478 EMW: T. Lincoln: “wolves could not kill “ moose even though the moose “were plenty enough”).

54. Historical Evidence of Two Species of Wolves in Maine continued 3) Only two wolves from Maine in museums. Both at Harvard University, and both tested genetically as red wolves. Both killed in the late 1800s after the grey wolf was apparently extirpated. 4)Today, coyotes in Maine show more genetic evidence of the red versus gray wolf, suggesting that the gray wolf was extirpated first. 5)Historical prey distribution: caribou and moose (i.e., large prey) northern, as is the gray wolf; deer (a medium sized prey species) southern in it’s distribution, as is the red wolf.

55. The Snowshoe Cat and the Snowshoe Hare