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The Status of Southeast Alaska's Endemic Mammalian Fauna: a Small Mammal Investigation Including the Prince of Wales Flying Squirrel

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The Status of Southeast Alaska's Endemic Mammalian Fauna: a Small Mammal Investigation Including the Prince of Wales Flying Squirrel The Status of Southeast Alaska's Endemic Mammalian Fauna: A Small Mammal Investigation including the Prince of Wales Flying Squirrel Final Report on RWO #31 of the NBS AKFWRU-Fairbanks submitted to: The US Fish and Wildlife Service Attn: John Lindell PO Box 021287 Juneau, AK 99802-1628 30 April1997 Joseph A. Cook, Christopher J. Conroy, and John R. Demboski Institute of Arctic Biology, University of Alaska Museum University of Alaska Fairbanks Fairbanks, AK 99775 Executive Summary This final report provides an overview of research to date on "The Status of Southeast Alaska's Endemic Mammalian Fauna: A Small Mammal Investigation including the Prince of Wales Flying Squirrel", a project funded through the US Fish and Wildlife Service, Southeast Alaska Ecological Services program. Although the Research Work Order was shonened to a single year due to a loss offunds, supplemental funding was received from the USDA Forest Service that allowed completion of some of the work. We are examining genetic differentiation in three mammalian species groups (flying squirrels, long-tailed voles, and red-backed voles) across the Alexander Archipelago and nearby mainland. These species groups include jour island endemic taxa (Microtus coronarius, Glaucomys sabrinus grisei(rons, Clethrionomys gapperi solus, and g. wrangeli) and another five taxa (£. gapperi stikinensis, C. g. phaeus, C. rutilus glacialis, Microtus longicaudus littoralis, Glaucomvs sabrinus zaphaeus) that are largely endemic to Southeast Alaska. All of these unique taxa need to be critically reviewed under the Endangered Species Act due to their limited distributions, occurrence in areas that have been subject to intensive forest management practices, and the relatively high "sensitivity" to future landscape disturbances (Swanston et al., 1996). In addition to interest in the conservation of these taxa, this project meets the objectives and primary recommendations of recent reviews of wildlife on the Tongass National Forest (Suring et al. 1992; Kiester and Eckhardt 1994; Yensen et al. in press; USDA 1991, 1996). To date, our investigations suggest that: 1) the mitochondrial cytochrome b gene provides insight into the genetic relationships of the island and mainland populations of the three taxa (i.e., this is an appropriate marker for these questions); 2) levels of genetic differentiation are relatively low for the Nonhern Flying Squirrel (Glaucomys sabrinus) across its range from Interior Alaska to Southeast Alaska; 3) that even within this low level of variation, the Prince of Wales Flying Squirrel (Glaucomys sabrinus grisei(rons) may be a valid taxon based on fixed nucleotide substitutions that identify all individuals of populations examined from Prince of Wales(15 individuals) and several islands located west of POW (6 islands; 23 individuals); 4) higher levels of genetic differentiation are found in the two long-tailed vole species [Microtus longicaudus (32 individuals) and M. coronarius (8 individuals)]; 5) this variation in individuals of Microtus identifies few unique populations or island groups, although the island populations appear distinct from most populations sampled on the mainland; and 6) Clethrionomvs rutilus and C. gapperi are not sister taxa, yet these species may show limited introgression near the mouth of the Stikine River. BACKGROUND The role of taxonomy and systematics in conservation biology, land management practices, and compliance with federal laws is well established (Avise 1994; Vane-Wright et al., 1991). Federal legislation (NEPA 1973, ESA 1973, NFMA 1976, TTRA 1990) has mandated the conservation of biological diversity. Considerable public and scientific attention is now focused on the loss of biodiversity on federal lands, particularly in landscapes that have been modified by human activity such as logging (e.g., Maser, 1994) . One of the main facets of maintaining biodiversity is the ability to recognize and conserve unique (endemic) taxa. More than 80% of Southeast Alaska (Figure 1) is included in the Tongass National Forest which manages some of the last significant old growth rainforest found in North America. Effective management within the multiple use approach, in addition to a reasonable understanding of ecosystem patterns and processes, requires baseline documentation of distribution and taxonomy of the fauna and flora. This information is missing for many mammalian taxa that are endemic to Southeast Alaska resulting in a serious gap in the ability of wildlife managers, biologists, and land planners to critically evaluate impacts of perturbations on potentially threatened or endangered taxa (Kiester and Eckhardt, 1994; Y ensen et al., in press). Attempting to manage a fauna for which little basic distribution and taxonomic data exists can lead to serious conservation problems (e.g., Daugherty et al., 1990; May, 1990). In Southeast Alaska, this situation is exacerbated by the highly insular and naturally fragmented nature of the landscape that results in an increased potential for biotic endemism. Indeed, 27 mammal taxa are endemic to Southeast Alaska; the largest number of endemic mammals of any US National Forest. Distribution and taxonomic status of the terrestrial mammal species 2 has been reviewed (MacDonald and Cook, 1996) and is, in general, poorly documented for most species. The need to characterize these taxa is urgent, given the rapid habitat modification that is occurring on islands with a high potential for endemism. Forest fragmentation in Southeast Alaska has resulted in extensive tracts of 100-year-old-or-less stands of closed-canopied forests; these second-growth forests generally support a less diverse mammal fauna (e.g. , Parker et al. , 1996). Islands in general are known for their high proportion of endemics (Quammen, 1996) due to the increased potential for divergence in isolation. The Tongass National Forest includes one of the largest island archipelagos in North America, yet, wildlife management plans and environmental assessments related to timber harvests have seldom discussed the potential impact of habitat manipulation on the endemic fauna. Insular forms are noteworthy from a conservation point of view as many animal extinctions observed in recent history have occurred on islands (Diamond, 1984; Soule, 1983). We have used molecular markers to begin to characterize the relationships, potential genetic divergence, and taxonomic status of several of the purported endemics of Southeast Alaska. Three distinct projects were supported in this study. These projects focused on flying squirrels, long-tailed voles, and red-backed vole populations across the Alexander Archipelago and nearby mainland. These species groups include four island endemic taxa (Microtus coronarius, Glaucomys sabrinus griseifrons, Clethrionomys gapperi solus, and C. g. wrangeli) and another five taxa (C. gapperi stikinensis, C. g. phaeus, C. rutilus glacialis, Microtus longicaudus littoralis, Glaucomys sabrinus zaphaeus) that are largely endemic to Southeast Alaska. Methods DNA extraction, amplification, and sequencing Frozen (-80°C) or alcohol preserved liver or heart samples from the Alaska Frozen Tissue Collection of the University of Alaska Museum (Table 1) were subjected to proteinase K digestion, NaCl precipitation of proteins, and DNA precipitation with ethanol, following a protocol modified from Miller et al. (1988). Partial or complete cytochrome b sequences (see individual projects) were amplified by the polymerase chain reaction (PCR; usually 30 cycles alternating denaturation at 93 oc for 1 min, annealing at 45 oc for 1 min, extension at 72 oc for 1.5 min) using combinations of the following primer pairs: MVZ05/MVZ04; MVZ14/MVZ23, and MVZ16/MVZ37 (Smith and Patton, 1993) and newly designed primers Cleth1, Cleth37 and Vole23, Vole14 (Table 2). Aliquots of the PCR products were visualized in 1.5% agarose minigels. The remaining product was precipitated with polyethylene glycol, recovered by vacuum centrifugation, and resuspended in 1X TE buffer. Purified products were used as template in cycle sequencing utilizing a Perkin-Elmer kit (Fst-RR, 402119) and run on an automatic sequencer (ABI 373). Both heavy and light strands were sequenced in all cases. Sequence and phylogenetic analysis Sequences were examined, assembled based on overlapping regions and aligned using the Sequence Navigator version 1.01 (Applied Biosystems Inc). Nucleotide composition and numbers of variable and phylogenetically informative sites at both the nucleotide and the amino acid levels (Table 3) were obtained using the program MEGA version 1.01 (Kumar et al., 1993). Phylogenetic analyses under the criterion of maximum parsimony were carried out using PAUP (version 3 .1.1, Swofford, 1993). Details of these analyses are described in the individual species accounts. 4 PROJECT I Genetic differentiation of the Prince of Wales Flying Squirrel, Glaucomys sabrinus griseifrons, an old-growth forest dependent mammal in Southeast Alaska John R. Demboski and Joseph A. Cook The northern flying squirrel, Glaucomys sabrinus, has a broad distribution across Interior Alaska, most of Canada, and south into the major mountain ranges of western and eastern North America (Hall, 1981). Two endemic subspecies of northern flying squirrels are known from Southeast Alaska. The Alaska coast flying squirrel, Glaucomys sabrinus zaphaeus, was originally described by Osgood ( 1905; type locality Helm Bay, Cleveland Peninsula). Later, Howell (1934) described the Prince of Wales Island,
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