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1 Future Population Trends and Drivers of Change for Alexander Archipelago Wolves on and Near Prince of Wales Island, Alaska

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1 Future Population Trends and Drivers of Change for Alexander Archipelago Wolves on and Near Prince of Wales Island, Alaska Gilbert et al. (2015) Final Report 4 Sep. 2015 FUTURE POPULATION TRENDS AND DRIVERS OF CHANGE FOR ALEXANDER ARCHIPELAGO WOLVES ON AND NEAR PRINCE OF WALES ISLAND, ALASKA Sophie L. Gilbert1, Trevor Haynes1, Mark S. Lindberg1,2, David M. Albert3, Michelle Kissling4 1Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, Alaska, 99775 2Department of Biology and Wildlife, University of Alaska Fairbanks, Fairbanks, Alaska, 99775 3The Nature Conservancy, Juneau, Alaska, 99801 4The U.S. Fish & Wildlife Service, Juneau, Alaska, 99801 Final Report for Cooperative Agreement: F15AC000206 September 4, 2015 1 Gilbert et al. (2015) Final Report 4 Sep. 2015 ABSTRACT The Alexander Archipelago wolf, inhabiting the coastal temperate rainforest of North America from British Columbia to southeastern Alaska, was recently petitioned for protection under the U.S. Endangered Species Act. Concerns about habitat alteration from industrial timber harvest and subsequent declines in prey (deer), increased human-caused mortality, and climate change were presented as the basis for the listing. To evaluate how these factors will likely affect future trends in abundance for Alexander Archipelago wolves and deer, we constructed a model linking wolf and deer population dynamics to environmental conditions and management regulations. We restricted our model to Prince of Wales and outlying islands, because this area is partially isolated, is the focus of timber harvest in the region, and has the most empirical data available for model parameterization. We examined the effects of timber harvest (past and future), winter severity, wolf harvest regulations, and roads, which affect deer and wolf harvest, on population dynamics of deer and wolves. Combining these factors, 6 future scenarios were developed by a panel of experts, and subsequently evaluated using the model. The ecological backdrop of this study is that high rates of logging during the 1970’s and 1980’s, and the lag in time over which habitat values for deer subsequently decline due to forest succession, resulted in a significant downward trajectory in carrying capacity for deer during 1995 – 2015, with varying levels of decline continuing through 2035 under all scenarios. Across scenarios, we found that wolf populations persisted, but with declines of 5−20% in wolf abundance and 21−32% in deer abundance after 30 years, with variation produced primarily by changes in road density, and smaller contributions from vegetation scenarios, severe-winter frequency, and wolf harvest regulation. In addition, we found that wolf declines could be greater if wolves rely more heavily on deer in the future, for instance if salmon availability declines under future climate change. 2 Gilbert et al. (2015) Final Report 4 Sep. 2015 Various wolf harvest regulations (0%, 20% and 30% caps on reported harvest of the fall wolf population) did not produce large differences in wolf abundance because the harvest caps were calculated at the population level, which were frequently not met with our modeling approach. The potential importance of unreported harvest in wolf population dynamics needs further acknowledgement and treatment. Changes in deer harvest produced large increases in deer and wolf abundance. Although we evaluated factors affecting wolf abundance individually, we encourage a holistic approach to management of this predator-prey system in an altered ecosystem. KEYWORDS: Alaska; U.S. Endangered Species Act; Alexander Archipelago wolf; predator- prey interaction; population dynamics INTRODUCTION The coastal temperate rainforest of the Pacific Northwest of North America is a globally rare ecosystem, with an assembly of species found nowhere else in the world. Within this eco-region, timber harvest and associated development have altered the landscape considerably over the last century, especially in the last 50 years, resulting in a patchwork of even-aged forest stands, intersecting roads, and small human settlements. The majority of remaining old-growth temperate rainforest in the U.S. is found in southeastern Alaska, the northernmost extent of the coastal temperate rainforest ecosystem, which is the focus of both the timber industry and conservation efforts. One of the most controversial species inhabiting the coastal temperate rainforest of southeastern Alaska is the Alexander Archipelago wolf (Canis lupus ligoni), although its 3 Gilbert et al. (2015) Final Report 4 Sep. 2015 subspecific designation currently is being debated (e.g., Chambers et al. 2012; Cronin et al. 2015; Weckworth et al. 2015). Morphologically, it is smaller and darker, with coarser and shorter hair compared to continental gray wolves (Goldman 1944, Wood 1990). Genetically, coastal wolves in southeastern Alaska and British Columbia are distinct from continental populations (Shields 1995; Weckworth et al. 2005; Weckworth et al. 2010; Munoz-Fuentes et al. 2009; but see Cronin et al. 2015). Within this coastal region, wolves exhibit some genetic population structure, especially on POW where the population is partially isolated (Weckworth et al. 2005, Breed 2007, Cronin et al. 2015b). The debate regarding taxonomic classification of the Alexander Archipelago wolf largely stems from the lack of a universally accepted definition for subspecies (Haig et al. 2006) and the absence of a comprehensive morphological and genetic assessment of coastal wolves. We assumed that the subspecies designation is valid and refer to coastal wolves as the Alexander Archipelago wolf throughout this report. This assumed subspecies of the gray wolf depends on Sitka black-tailed deer (Odocoileus hemionus sitkensis) as a primary food source, and is consequently affected by timber harvest as deer populations respond to post-harvest forest succession (Figure 1). This trophic link from wolves to deer to timber harvest, along with high harvest of wolves in in areas accessible by road and boat (Person and Russell 2008, Person and Logan 2012), has resulted in concern for the long-term viability of wolves in southeastern Alaska. A major population of Alexander Archipelago wolves, inhabiting Prince of Wales and outlying islands, has declined sharply in recent years (Alaska Department of Fish and Game 2015a), further heightening concern. In 1993, a petition was filed for protection of the Alexander Archipelago wolf under the Endangered Species Act (ESA), but ultimately the U.S. Fish and Wildlife Service (USFWS) determined that listing as threatened or endangered was not warranted (62 Federal Register 46710, September 4, 4 Gilbert et al. (2015) Final Report 4 Sep. 2015 1997). In 2011, a petition was again filed for protection of the Alexander Archipelago wolf under the ESA, citing cascading effects of past and future timber harvest, high rates of wolf harvest, climate change, and inadequacy of existing regulatory mechanisms as primary threats (Center for Biological Diversity and Greenpeace 2011). The USFWS published a positive 90-day finding in the Federal Register in March 2014, indicating that the petition included substantial information suggesting that listing may be warranted (79 Federal Register 17993, March 31, 2014), and initiated a status review shortly thereafter. Per a court agreement, the USFWS must issue a 12- month finding on whether or not listing is warranted on or before December 31, 2015. As part of this process, we were charged with conducting a population analysis of wolves on Prince of Wales Island (POW), the largest island in the Alexander Archipelago, along with associated smaller islands, and this final report summarizes our results. Like their continental counterparts, coastal wolves are opportunistic predators, although their diet includes a greater marine component. Wolves in the coastal temperate rainforest prey on diverse species including Pacific salmon (Onchorhynchus spp.), mountain goat (Oreamnos americanus), marine mammals (e.g., harbor seal [Phoca vitulina]), American beaver (Castor canadensis), and a variety of birds. Although the diet of coastal wolves is diverse, deer is the most important prey item. In southeastern Alaska, deer typically represent at least half of annual wolf diet and Pacific salmon account for upwards of 20% of the annual wolf diet (Szepanski et al. 1999, Person 2001), although during late summer and autumn when adult salmon enter rivers to spawn, salmon can be the dominant prey item (Darimont et al. 2008). The proportion of deer in wolf diets varies considerably by population, by pack, and among individuals, although much of the variation among populations is explained by differences in availability of deer and alternative large-bodied mammalian prey (Darimont et al. 2004). As a result, wolves in island 5 Gilbert et al. (2015) Final Report 4 Sep. 2015 populations rely more heavily on deer than in mainland populations, where other large mammalian herbivores are available and deer are less abundant (Szepanski et al. 1999, Darimont et al. 2004). Therefore, on islands like POW, wolves are linked strongly to deer, although salmon subsidize wolf diets during parts of the year, thereby reducing dependency on deer to some extent (Szepanski et al. 1999). The primary drivers of deer population dynamics in southeastern Alaska include winter severity, habitat quality, and predation by wild carnivores and humans. Winter severity is highly variable through time and space in southeastern Alaska and coastal British Columbia (Shanley et al. 2015), and plays a key role in governing deer survival and population dynamics (Person
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