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Monitoring Recovery of Overgrazed Lichen Communities on Hagemeister Island, Southwestern Alaska Patrick Walsh1 & Trevor Goward2

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Monitoring Recovery of Overgrazed Lichen Communities on Hagemeister Island, Southwestern Alaska Patrick Walsh1 & Trevor Goward2 Monitoring recovery of overgrazed lichen communities on Hagemeister Island, southwestern Alaska Patrick Walsh1 & Trevor Goward2 1 U.S. Fish and Wildlife Service, Togiak National Wildlife Refuge, P.O. Box 270, Dillingham, Alaska 99576, USA 1 (Corresponding author: [email protected]). 2 UBC Herbarium, Beaty Museum, University of British Columbia, Vancouver, BC V6T 1Z4, Canada. Abstract: Understanding the recovery rate of overgrazed lichen communities has value to mangers of lands in northern regions. We describe lichen community composition and present recovery rate measurements for a 12-year period following overgrazing by reindeer (Rangifer tarandus) on Hagemeister Island, Alaska. Reindeer were removed from the island in 1993 following overgrazing and average total lichen biomass increased from 504.2 kg/ha (SD 205.4) in 2003 to 795.3 (SD 489.6) in 2015. We estimate time to recovery with three competing growth curves which estimate grazeable biomass may be reached in 34-41 years. However, estimates of full recovery to climax biomass varied among the models, ranging from 71 to 400 years. In 2015, lichen communities were composed of various mixtures of at least 78 lichen taxa, and were dominated by Cladina stygia and other important reindeer forage species. While reindeer overgrazing diminished forage quantity, it did not extirpate preferred forage taxa. Key words: Range recovery; Rangifer tarandus; grazing management. Rangifer, 41, (1), 2021: 1-12 DOI 10.7557/2.41.1.5340 Introduction Hagemeister Island, located in Bristol Bay to reduce the reindeer population from 1,000 to of the Bering Sea, is characterized by various 450 (Swanson & LaPlant, 1987). The reindeer shrub and tundra plant and lichen communi- population was not reduced, and a 1987 survey ties (Fig. 1, Swanson & LaPlant, 1987). These found damage to lichen sites but little impact communities developed in the absence of un- to vascular plant-dominated sites (Swanson & gulate grazers until 1965, at which time rein- LaPlant, 1987). In 1992, Swanson and Barker deer were introduced. The reindeer population (1993) reported virtual depletion of lichens on grew from an initial introduction of 71 ani- the primary lichen-dominated sites and signifi- mals to over 1,000. Overgrazing was evident cant ecological damage on the remaining vascu- by 1973, at which time a recommendation was lar-plant dominated sites. A die-off of approxi- made to the U.S. Bureau of Land Management mately 300 reindeer occurred in 1992 and the This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Rangifer, 41, (1) 2021 Editor in Chief: Birgitta Åhman, Technical Editor: Eva Wiklund and Graphic Design: H-G Olofsson, www.rangiferjournal.com 1 remaining animals were removed in 1992-1993 of that on Hall Island, suggesting that in this by the U.S. Fish and Wildlife Service (which had example, post-grazing recovery may require a been delegated management authority in 1980) similar amount of time as that following fire. in order to end the ecological effects caused by Swanson estimated that it would take 75 – 100 overgrazing (Swanson & Barker, 1993). years for the depleted lichen communities on Lichens are important in reindeer and cari- Hagemeister Island to recover in the complete bou (both R. tarandus) diets, particularly dur- absence of reindeer grazing (letter to Togiak ing winter (Klein, 1968; Pegau, 1968; Skoog, National Wildlife Refuge manager in Swanson 1968; Thompson et al., 1981; Boertje, 1984; and Barker, 1993). However, other investiga- Joly et al., 2007; Joly & Cameron, 2018). Li- tors have suggested that recovery following chen taxa that have been determined to be par- overgrazing can occur more rapidly. Henry ticularly important forage species to reindeer and Gunn (1991) reported that essentially all and caribou include members of the principally macrolichen biomass had been eliminated in ground-dwelling genera Cladonia, Cladina, Ce- a single summer grazing event on an island in traria and Stereocaulon and the arboreal genera Northwest Territories and suggested that recov- Alectoria, Evernia, and Usnea (Skoog, 1968; ery could take 20 years or longer. Collins et al., Sveinbjornsson, 1990). Lichens are character- (2011) evaluated the impacts of caribou graz- ized by extremely low rates of growth relative to ing on lichen abundance and subsequent use in vascular plants (Pegau, 1968; Sveinbjornsson, interior Alaska, reported that caribou selected 1990). Lichen abundance can be reduced by for habitat with >1,250 kg/ha forage lichen overgrazing, as in the Hagemeister Island case biomass, and hypothesized that substantial re- or by fire, which tends to be a more destruc- covery following overgrazing may only require tive process generally requiring longer recovery 20 years. Our goals are to document the rate of times than that required by overgrazing, given recovery of lichens on Hagemeister Island fol- that recovery post-fire is often dependent upon lowing the removal of reindeer and present al- primary succession over large areas (Collins ternative recovery scenarios, determine the rela- et al., 2011). There have been numerous in- tionship between biomass and visual estimates vestigations of lichen recovery post-burning, of lichen cover, and describe lichen community given that much of the range of caribou occurs composition. in the boreal north where fire is a regular eco- system process (Collins et al., 2011; Black & Study area Bliss, 1978; Kumpula et al., 2000; Maikawa Hagemeister Island (58o 29’ N, 160o 54’ W) is & Kershaw, 1976; Morneau & Payette, 1989; approximately 39 km in length, approximately Scotter, 1964). These studies are in agreement 31,200 ha in area, and is located approximately that recovery takes 100 years or longer. Klein 31 km south of the village of Togiak in Bris- (1968 and 1987) and Klein and Shulski (2009) tol Bay (Fig. 1, Bailey, 1983). The climate on studied lichen community recovery on St. Mat- Hagemeister Island is subarctic maritime. An- thew Island, Alaska following overgrazing by nual precipitation averages 94 cm and tempera- reindeer that was followed by a complete die- tures range from average daily low and high of off. Twenty-two years after the die-off, lichens –11 and –7 o C in February, the coldest month, had recovered to only 6% of the standing crop to 8 and 12o C in August, the warmest month at of living lichen biomass occurring on adjacent the nearest weather station, Cape Newenham, Hall Island with no grazing. At 41 years af- Alaska (climate data averaged 1981-2010, Na- ter the die-off, lichens had recovered to 12% tional Oceanic and Atmospheric Administra- This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License 2 Editor in Chief: Birgitta Åhman, Technical Editor: Eva Wiklund and Graphic Design: H-G Olofsson, www.rangiferjournal.com Rangifer, 41, (1) 2021 tion, 2019). The island surficial geology ranges training or other efforts were made to attempt from relatively recent flood plain alluvium, col- to standardize estimates. We then plucked all luvial deposits, and glacial drift of sand, gravel, fruticose and foliose lichen from each sam- and boulders to Paleozoic siltstone, limestone, pling frame and did not separate living from and argillaceous rocks (Hoare & Coonrad, non-living lichen. We oven dried the samples 1961). Intermediate in age are extensive car- at 40o C, sorted them into like taxa and then boniferous to cretaceous volcanic deposits. The weighed them to the nearest 0.001 g using an island ranges from rolling hills up to 150 m el- AND GX 400® balance (mass range: 0.001g evation in the east to steep mountains up to to 410g). All lichen identifications were per- 500 m in the west. formed by T. Goward using a dissecting micro- Hagemeister Island is treeless, composed of scope and chemical reagents as needed (Brodo 12 landcover types, and is dominated by vari- et al., 2001). ous shrub types ranging from tall to dwarf shrub We totaled lichen biomass of the 4 sub- (Collins, 2003; Winfree, 2009). Low eleva- samples taken at each site and considered the tion sites beginning at the beaches are forb and total biomass by taxa for each site as replicates. graminoid-dominated, and graminoid-dom- We tested for change in biomass and cover be- inated communities also occur above coastal tween sampling periods using unequal-variance cliffs and on elevated benches. Extensive alder T-tests when Kolmogorov-Smirnov tests in- (Alnus) thickets occur on mountain slopes pro- dicated data were not normally distributed or tected from the wind. Lichens occur primar- demonstrated unequal variance. When nor- ily on exposed upland dwarf shrub lichen sites, mally distributed, we made comparisons with which by definition include >20% lichen cover equal-variance T-tests. Estimated lichen cover (Viereck, 1992). Hagemeister Island supports was averaged among plots at each site and sites diverse faunal communities, but in the absence were considered replicates. Average cover val- of reindeer includes no lichen grazers (Bailey, ues were related to total biomass of all lichens 1983; Stimmelmayr, 1994). per site using ordinary least squares regression. Statistical tests were performed at an alpha level Methods of 0.05 using NCSS 10 Statistical Software We randomly selected 27 study sites from (2015). dwarf shrub lichen sites >4 ha to minimize the We calculated the rate of lichen recovery us- probability of sampling in the ecotone of other ing 3 different models: linear and exponential cover types (Fig. 1). At each site, we collected 4 growth functions and a growth curve specifical- samples at 3.05-m intervals along a randomly- ly developed for woodland lichens in Scandi- oriented transect originating at the approximate navia (Pekkarinen et al., 2015). Linear growth sample site center. Samples consisted of lichens was calculated as the average yearly increase occurring in 0.125 m2 rectangular sampling in average lichen biomass between 2 points in frames.
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