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Livestock in Alaska | Cooperative Extension Service | Cooperative Extension Service
Certain persons may require specific supporting documentation such as an employment petition, student authorization, or approval notice. For more details, go to the website. Foreign visitors entering the U. The northern shift in the distribution of moose, like that of snowshoe hares, has been in response to the spread of their shrub habitat in the Arctic, but at the same time, herbivores have likely had pronounced impacts on the structure and function of these shrub communities.
These northward range shifts are a bellwether for other boreal species and their associated predators. This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication. Vegetation and environmental gradients of the Prudhoe Bay region, Alaska.
Report no. All other relevant data are within the paper and its Supporting Information files. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist. Temperatures in the Arctic increased rapidly during the 20 th century following centuries of cooling [ 1 , 2 ], and the resulting landscape changes, including increased vegetation productivity and the expansion of shrubs, have been widespread [ 3 — 5 ].
The effects of warming and landscape changes on tundra wildlife, in comparison, are poorly documented and limited in spatial or temporal extent. Arctic vegetation in Alaska has been altered by 20 th century warming, yet little affected by direct human impacts, so the region provides a setting to examine the effects of altered habitat on wildlife. Here, we review historical sources across northern Alaska and the pan-Arctic to examine over a century of climatic influence on shrub habitat and moose Alces alces.
History of Alaska
We focus our study on moose in the Alaskan tundra because their shrub habitat is known to have increased, and because their large size, unmistakable appearance, and importance as a source of protein for people lent them to historical documentation. In northern and western tundra regions of Alaska, the lack of moose during the 19 th and early 20 th century was tentatively attributed to hunting by indigenous peoples and miners, and the subsequent expansion of moose has been associated with human emigration from inland regions and the resulting reduction in hunting [ 6 , 7 ].
To evaluate whether current moose presence in the Alaskan tundra is due to 20 th century warming and expanded shrub habitat, we used the change in cumulative summer warmth thaw degree days from to , combined with empirical correlations between thaw degree days and shrub height, to estimate riparian shrub height starting in We compare these reconstructions of shrub height to the habitat preferences for moose [ 12 ] to evaluate whether suitable habitat existed during periods when moose were absent from the Alaskan tundra.
We discuss shrub habitat expansion alongside other potential factors, such as hunting and wolf predation, in facilitating moose expansion into tundra regions. Moose are the largest member of Cervidae and occupy a diversity of north temperate ecosystems. Although predation, disease, and weather influence population dynamics, suitable climate and habitat facilitate the establishment and persistence of populations, thereby shaping the regional distribution of moose [ 13 ].
The northern edge of moose distribution generally follows latitudinal treeline, occasionally extending northward into tundra along major riparian corridors [ 14 , 15 ]. Moose are mainly associated with early-successional habitats and selectively feed on relatively high-quality riparian shrubs, particularly willow Salix spp. In tundra and ecotonal regions, availability of forage shrubs above the snow is limiting [ 16 , 17 ].
Moose also utilize dense woody vegetation as cover to reduce detection by predators [ 18 , 19 ], and as protection during wolf attacks [ 20 , 21 ]. Longer winters and correspondingly shorter shrubs coastward [ 22 ] reduce habitat suitability and likely explain why moose distribution ends some distance south of the coast in tundra regions of Alaska, Canada, and Siberia. The assemblage of carbon dated moose bones suggests at least a periodic presence of moose in the region during the last three millennia, though the date range on some of the more recent bones indicate that they could be modern.
Evidence from archeological sites, indigenous peoples, and early explorers documents an absence of moose during the latter half of the 19 th and early 20 th century in tundra areas of Alaska [ 6 , 24 — 26 ], with few exceptions [ 6 , 27 ]. During the second quarter of the 20 th century, moose began to appear in tundra regions of northern and western Alaska Fig 1 , and similar increases were observed in northern Canada A.
By the s, moose populations were becoming established along the riparian shrub corridors of the Colville River and its tributaries in Arctic Alaska. An aerial population survey in late-winter of revealed moose along a km section of the Colville River floodplain [ 6 ].
Extensive late-winter surveys in and covering most of the North Slope Utukok to Kongakut River recorded between and moose [ 6 ], with approximately half of those moose residing in the middle Colville drainage [ 30 ], confirming their establishment. By the s moose had colonized northwest Alaska Fig 1 [ 31 ]. Map is inset from Fig 3. Shrub plots were distributed along the Chandler and Colville Rivers orange ellipse , and temperature records were derived at two locations therein gray dots [ 33 ].
We used the relationship between summer warmth and willow height developed by Walker [ 22 ] to estimate changes in the height of riparian willows from c. Walker [ 22 , 32 ] measured the 50 tallest willows Salix richardsonii at multiple streamside sites along a temperature gradient in Arctic Alaska and derived a relationship between thaw degree days TDD and shrub height: Eq.
We assumed that shrub height was a function of the past 10 years of summer warmth, and therefore used an average of the previous 10 years of thaw degree days to estimate shrub heights annually since In , we measured heights of streamside shrubs, including the 50 tallest willows Salix spp. These two riparian corridors were selected because they have the greatest density of tall shrubs and moose north of the Continental Divide of the Brooks Range [ 30 , 37 ], and would likely have been the first riparian corridors with sufficient habitat for moose.
We compared our predictions of shrub heights in from Eq 1 to our measured values in Mean and standard error are reported, unless otherwise mentioned. Our hindcasting of shrub height, based on the strong positive relationship between streamside willow height and thaw degree days [ 22 ], indicates that the shorter and cooler growing seasons c. The 1. Estimated changes in moose habitat in Arctic Alaska since , including changes in riparian shrub height a, b and shrub canopy volume c, d.
Moose require shrubs protruding above the snow in late winter for habitat [b; adapted from [ 12 ]], and the shrub height hindcasting a indicates that little to no habitat would have existed in Arctic Alaska c. The blue shrub height line uses interpolated observed temperatures, whereas the red line uses temperatures hindcasted from an average of the five highest-performing General Circulation Models [ 34 ]. The mean July temperatures of the late s and early s solid line is average of previous 10 yrs were less than Temperature sensitivity of shrub canopy volume for Salix richardsonii , the species used to construct a , is similar to that for the preferred forage species of moose, Salix alaxensis d; reproduced from [ 36 ].
The spatial expansion and vertical growth of riparian shrubs in response to warming initiated in the 19 th century would have substantially increased winter moose habitat. The estimated tall shrub height of 1. Salix richardsonii , the willow species measured along a thermal gradient [ 22 , 32 ] and used here, responds very similarly to mean July temperature exceeding Mean July temperature was below General agreement between the observed riparian shrub heights in 2. In tundra regions moose require shrubs protruding above the snow [ 16 , 17 , 38 ].
In riparian corridors, the valley topography and shrubs dampen the erosive wind events that commonly scour snow from the surrounding tundra, leading to snow depths more than twice as great in the riparian shrubs as on the surrounding tundra [ 39 ]. The mean late-winter snow depths of 0.
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Little available forage would have protruded above the snow prior to moose establishment, and an increase in average shrub height from 1. The increase in shrub height might have captured more drifting snow, potentially negating some of the gains in available forage, but the scant record of historical snow distribution led us to assume a variable but trendless end-of-winter snow cover amid increasing shrub heights.
Finally, adult moose in Alaska stand approximately 1. The only proxy record of shrub production dating to the 19 th century was reconstructed from sediment cores collected from the Colville River delta, and it shows a much greater increase in shrub production as indicated by increases in fresh particulate organic matter transported to the delta in the Colville watershed between and than after [ 8 ], when repeat photography documents an increase [ 41 ].
Shrub expansion initiated c. Summer warming prior to is evident from historic photos showing the retreat of glacier terminuses in Arctic Alaska [ 42 ] from their Little Ice Age maximum extents—retreat that had accelerated by [ 43 ] and continues today [ 44 ]. Increasing mean annual air temperatures 50 to 75 years leading up to the s was also evident in warming permafrost borehole temperature profiles from the region [ 45 ]. Warming inferred from proxy records is consistent with temperature data generated by running GCMs backward in time, which show an increase in summer temperature between and Fig 2A and 2C.
Our result of increasing shrub habitat linked to moose establishment can confidently be extrapolated across the North Slope and Brooks Range, where summer temperatures are comparable and there is a record of shrub habitat increase.