Novel evidence that elk were historically native to the Sierra Nevada, and recent range expansions into the region

Elk (Cervus canadensis) have been considered non-native to the Sierra Nevada mountain range of California and Nevada. However, elk have steadily increased their range southward from the Cascade Range into the northern Sierra Nevada over the last century. Recent reports also reveal Rocky Mountain elk moving northwards into the southern Sierra Nevada. Dispersals of lone bull elk from 2019–2022 have occurred to the central Sierra Nevada south of Lake Tahoe. These recent range expansions of elk herds and long-distance dispersals of individual elk raised questions about the possible historical presence of elk throughout this mountain range. Herein we conducted a broad investigation into historical newspaper accounts and other early explorer and naturalist observer records, museum specimens, Late Holocene zooarchaeological records, and indirect evidence including toponomastic references and Native American ethnographic and ethnolinguistic information. Taken in total, a variety of data sources suggest elk inhabited portions of the Sierra Nevada and the adjacent northwest Great Basin from the Late Holocene through historical times. Positive records were not numerous, suggesting that historically elk were not abundant, and nearly extirpated during the California Fur Rush of the early nineteenth century.

Yes -all data are fully available without restriction  Range" (Fig. 1).Tule elk have a well-established habitat preference for more open terrain and generally avoid forested habitat and higher elevations characteristic of the Sierra Nevada (3).This strong habitat preference is likely related to physical and behavioral adaptations for drier, Mediterranean-climate conditions (4,5).In describing primitive North American elk distribution in 1951, O.J. Murie cited a lack of available records of elk in easternmost and southernmost California and suggested that elk did not occupy Nevada "in any considerable numbers".Murie recognized, however, that elk have occurred beyond the boundaries presented and a more extensive record would likely reveal a greater historical distribution (2).In 1969 D.R.

McCullough identified reliable historical records and museum specimens consistent with Rocky
Mountain elk historical presence in Shasta and Siskiyou Counties of northeastern California (3,6,7).Until then California Fish and Game had disputed that this subspecies had existed in California, aligning with Murie's perspective (1,8).McCullough's findings were accepted by CDFW in their 2018 elk plan (1).In partial agreement with Murie's work, the 1997 Nevada Department of Wildlife (NDOW) Elk Species Management Plan established that elk were native to Nevada, but densities were likely kept low by Native American hunters wherever they occurred (9).In contrast to the CDFW Elk Management Plan, the NDOW Elk Species Management Plan noted that "Newspaper accounts report hunter kills at Lake Tahoe and Honey Lake Valley…" (10).These reports, both near the California and Nevada border in the Sierra Nevada region, were based on an extensive search of historical newspaper and other observer records, but the specific citations were not listed.Nevertheless, these reports raise questions about the historical presence of elk in the Sierra Nevada region.
Few contemporary studies are conducted with their primary goal the determination of the historical range of a given species (11).Understanding ecosystem reference conditions in California and Nevada present challenges due to the relatively late establishment of zoological collections in museums in these states until the late nineteenth and early twentieth century (12).
The California Fur Rush of the first half of the nineteenth century began as fur trappers and hunters ventured into the Sierra Nevada region beginning with Jedediah Smith in 1827 and Peter Skene Ogden in 1828 (8,13,14).These early venturers, and/or aboriginal hunters aided by the introduction of horses and firearms, may have rapidly depleted elk in the Sierra Nevada before early naturalists could document their presence.For example, after killing over 130 elk in a single winter season at the mouth of the Columbia River from November 1805 to March 1806, Lewis and Clark lamented how difficult it had become to hunt elk without traveling long distances from their camp (15).These factors may have led to similar underestimation of the historical ranges of elk and other mammalian species in California.In fact, several recent historical ecology studies have expanded the known historical range of species long thought to be non-native to the Sierra Nevada region, including physical evidence of the historical presence of North American beaver (Castor canadensis) in the high Sierra Nevada (12,16) and the historical nativity of the California red-legged frog (Rana draytonii) to the Yosemite Region (17).Although direct evidence of historical presence of the gray wolf (Canis lupus) in the Sierra Nevada has not been found (although one museum specimen was collected near Litchfield, Lassen County, California just north of the Susan River) (18,19), wolves have recently colonized both the northern and central Sierra Nevada, with packs currently (re-)established in Plumas and Tulare Counties, respectivelysuggesting that at least portions of the Sierra Nevada remains suitable wolf habitat (20).Similarly, an historical ecology study of the presence of wolverine (Gulo gulo) found historical observer records only in the southern subalpine and alpine Sierra Nevada (21), but recent discoveries of wolverine dispersers to the Lake Tahoe and Yosemite National Park regions in 2008 and 2023 raise questions about the former distribution of this carnivore in the Sierra Nevada (22).
The Sierra Nevada is dominated by coniferous forest, a habitat type often associated with Rocky Mountain elk, and there are no obvious biogeographical barriers that would prevent Rocky Mountain elk range expansion south from the southern Cascade Range to the northern Sierra Nevada.The uncited historical newspaper accounts mentioned above and recent studies expanding the historical range of other mammals to the Sierra Nevada, prompted our search for historical evidence of elk in the region.To overcome the paucity of evidence typical of historical ecology studies (23), we evaluated multiple different lines of evidence, similar to a recent investigation for evidence of late Holocene and historical records of bison (Bison bison) in northeastern California and northwestern Nevada (24).Our study goals were to research and unify broad data sources for historical and late Holocene evidence of elk in the Sierra Nevada, and to update information on the region's current (re)colonization by elk.

Study areas and time period
The Sierra Nevada mountain range is 640 km (400 miles) long on its north-south axis, and 80-130km (50-80 miles)) wide east-west.Its northern border stretches from the North Fork Feather River east to Fredonyer Pass south of Mount Lassen, then further east to the Susan River.
Its southern border is at Tehachapi Pass where the Sierra Nevada give way to the Tehachapi Mountains to the southwest (12).The Sierra Nevada's peaks steadily rise in elevation from 2,130 m in the north to 4,413 m tall Mount Whitney in the south.With the exception of zooarchaeological specimens dating to the Subatlantic age (< 2,500 years BP) of the Late or Meghalayan Holocene (25), the time period of the evidence investigated to support the historic presence of elk in California beginning with the arrival of Spanish explorers in 1769 (26).

Specimen and records search
First, we searched museum records and zooarchaeological specimens for physical evidence of elk's historical presence in the Sierra Nevada region.For museum specimens we examined the Arctos Multi-Institution and Multi-Collection Museum Database (Arctos) (https://arctos.database.museum/),the Mammal Networked Information System (MaNIS) (http://manisnet.org/),and the Integrated Digitized Biocollections (iDigBio) (https://www.idigbio.org)via Boolean searches using the terms "elk", "wapiti", "Cervus canadensis", and "Cervus elaphus".In addition, we contacted curators of mammal collections at the California Academy of Sciences (CAS), the Berkeley Museum of Vertebrate Zoology (MVZ), and the National Museum of Natural History (USNM) for elk specimens collected in the Sierra Nevada region.For zooarchaeological records of elk in the region we queried FAUNMAP (http://www.ucmp.berkeley.edu/neomap/search.html)for "Cervus elaphus" and "Cervus canadensis" remains.and references listed in recent reviews of large ungulates (bison and elk) in the Great Basin (24,27).
We used Web of Knowledge and Google Scholar to search for historical observer records by the earliest explorers and naturalists to the Sierra Nevada as well as those in citations in publications that reviewed the historical ranges of other California mammals in the region (12,19,21,(28)(29)(30).

Indirect evidence search
Indirect evidence searches included searches for geographic place names using the United States Geological Survey (USGS) Geographic Names Information System (GNIS) (https://geonames.usgs.gov/pls/gnispublic)and toponomastic references for California, Nevada, and the Lake Tahoe area (31)(32)(33).Because ethnographic and ethnolinguistic information does not provide a specific written historical record, we considered it indirect evidence.We researched Native American ethnographic and ethnolinguistic information and geographic place names including the word "elk" as referenced below.

Historical newspaper accounts and other observer records
First, we located the previously uncited references for the two historical newspaper accounts in the NDOW report.The first historical newspaper account was printed in The Carson Daily Appeal in 1867 (34) and reads as follows: "ELK AT LAKE BIGLER-We learn that two elk were killed at Lake Bigler, beyond the Zephyr Cove House, during last week.One of them weighed about five hundred pounds."-The Carson City Daily Appeal 1867.Lake Bigler was the historical name of Lake Tahoe, named for California's third governor, John Bigler.However, because Bigler was a secessionist, the United States Department of the Interior renamed it Lake Tahoe during the Civil War (32).Zephyr Cove House was established by Andrew Gardenier, who homesteaded 65 hectares (160 acres) and built the inn in 1862 on beachfront property in Zephyr Cove, in Douglas County, Nevada on Lake Tahoe's southeastern shores, to cater to silver miners (35).The second newspaper record is of an elk hunt in the vicinity of Honey Lake Valley, Lassen County, California (36):

"RETURN OF THE HUNTERS-About two weeks since Alderman Dimock and one or two other gentlemen left this city [Virginia City] for a hunt in the direction of Honey Lake… Night before last they returned to this city, bringing several trophies of their prowess. During their excursion they killed an elk, weighting 404 pounds, and a number of deer, antelope, and smaller game." -Territorial Enterprise 1868.
Honey Lake Valley is south of the Susan River, near the extreme northeastern edge of the Sierra Nevada, located 107 km north of Lake Tahoe.This account was printed in the Territorial Enterprise (Virginia City) on February 12, 1868 (Territorial Enterprise 1868).Both Honey Lake Valley and Lake Tahoe are south of the Susan River, placing both newspaper accounts in the Sierra Nevada (Fig. 2).The two newspaper accounts are 122 km and 260 km south, respectively of CDFW recognized historical elk range (Modoc County's southern border) (1).

Fig 2. Map depicting locations of nineteenth century elk hunts "H" at Honey Lake and
Lake Tahoe in the Sierra Nevada.Supporting evidence includes two Late Holocene zooarchaeological records of elk "Z", and suitability of habitat reflected by presence of elk today in the northern Sierra Nevada north of Lake Tahoe and the southern Sierra Nevada about Lake Isabella.
Despite a comprehensive search, we located a single early historical observer record of elk in the Sierra Nevada region, given by Thomas Jefferson Farnham, who explored California in 1840: "On the northwest side of this Desert [Great Basin] is a partially fruitful region, called the Vale of Mary's River [Humboldt River]… There are, however, some pretty groves of aspen and pine to be found along the stream and in the hills, among which live a few red deer and elk." (29).Farnham also noted the occurrence of elk even further south than the Sierra Nevada in California.In 1840 he described the course of the Mojave River as it approached its headwaters in the San Bernardino Mountains, as "gurgling through narrow vales covered with grass and fields and forests in which live the deer, the black bear, the elk, the hare, and many a singing bird" (29).The Mojave River mainstem begins in the San Bernardino Mountains, 128 km southwest of Tehachapi Pass, the southern border of the Sierra Nevada.John Boyden "Grizzly" Adams, resided at Long Barn in the Sierra Nevada, near Sonora Pass in Tuolumne County, and described hunting elk in the 1850's but these accounts were not clearly in the mountains versus the Central Valley of California and further may have been sensationalized for the San Francisco newspapers (37).
There are numerous place names including the word "elk" in California and Nevada, but only a single occurrence in the Sierra Nevada counties of these two states, an Elk Point in Douglas County, Nevada on Lake Tahoe (32,40).Coincidentally, Elk Point is only 0.8 km south of Zephyr Cove, although its etymology is unknown (32).Searches of other toponomastic references yielded no further elk place names in the Sierra Nevada (31,33).
In support of the historical newspaper accounts, there is ethnographic support for elk in the Sierra Nevada.The Washoe People had a word for elk, hañakmuwe (41).The Northern Paiute had a word for elk, but this does not provide added support since the Northern Paiute territory included portions of Oregon and Idaho where elk were historically present.However, there is a specific reference to the Honey Lake Valley Paiute, which were local to the Honey Lake Valley, stating that elk were "among the large animals captured" (42).Similarly, the Mountain Maidu's territory likely extended at least to the western edge of the Honey Lake Valley, and although the other side of their territory extended to the edge of the Central Valley, a fairly specific reference to elk as a food item in the mountainous region states: "In the mountains, deer, elk, mountainsheep, and bear were plenty; while in the Sacramento Valley there were great herds of antelope" (43).There is an Owens Valley Paiute legend of Hai'nanū who travels with his brother to Convict Lake then Mammoth in Mono County, California then over a ridge and shoots "soikwoi" or elk (44).

Reports of recent elk range expansions into the Sierra Nevada
The CDFW Elk Management Plan's current elk range maps indicate that by 2017, natural range expansion of Rocky Mountain elk had occurred southwards from Modoc County through Lassen, Plumas, and Sierra Counties, the latter 50 km north of Lake Tahoe (1).Elk calving grounds were identified in a 2020 study of the Plumas National Forest, which extends south to northern Sierra County, and reported that elk herds had been established there by the early 2000s (45).Presently, three spatially distinct herds occupy Plumas and Sierra counties, near Humbug reasonable specificity.A "500 pound" (227 kg) elk would not be a mule deer (Odocoileus hemionus hemionus), where adult bucks average 74 kg (163 lbs) dressed and about 25% heavier or 106 kg (233 lbs) live (48,49).Although mule deer size is quite variable, the largest trophy animal taken in northeastern California had a live weight of 181 kg (400 lbs) (50).The other newspaper account described a "404 pound" (184 kg) elk but differentiated it from deer and pronghorn (Antilocapra americana).The Honey Lake Valley newspaper account was supported by two nearby Late Holocene zooarchaeological records, one in California and one in western Nevada.This direct evidence is supported by a single reliable historical observer record near the eastern border of the Sierra Nevada by an early California explorer and naturalist, Thomas Farnham (29).
Indirect evidence also supports the historical presence of elk in the Sierra Nevada.
Ethnolinguistic and ethnographic evidence from four Native American tribes place elk in the Sierra Nevada and northwestern Great Basin historically.The toponomastic reference to an Elk Point, just south of the hunting record at Zephyr Cove on the southeastern shore of Lake Tahoe, while coincidentally close to the elk hunt there, may have an eponymous or other etymology unrelated to elk, the animal.Perhaps the strongest indirect evidence for the historical presence of elk in the Sierra Nevada is habitat suitability.Habitat in the northern Sierra Nevada is clearly suitable for Rocky Mountain elk as established by steady range expansions southwards in the northern Sierra Nevada.Further expansion of the nascent Lake Isabella herd in Kern County would suggest that the southern Sierra Nevada is also suitable Rocky Mountain elk habitat.
The paucity of historical observer records and Late Holocene zooarchaeological records are consistent with elk likely occupying the Sierra Nevada at very low densities historically.An extensive review of zooarchaeological records posited that elk were never abundant in the Great Basin, finding "20 Holocene-aged Great Basin sites positive for elk, compared to 77 that contained bison (Bison bison)-itself never an abundant animal in this region."(27).The lack of museum records is likely related to the relatively late establishment of museum zoology collections in California, which occur in the late nineteenth century only at the San Francisco California of Sciences and Stanford University's Museum of Natural History-after tule elk had been nearly extirpated (12).The other factor which may drive the rarity of museum specimens and historical observer records is that the Sierra Nevada was thinly settled and poorly explored by Anglo-Americans until the latter half of the nineteenth century.Similarly, late Holocene zooarchaeological specimens may have been rare, because Late Holocene Native American populations were low, so the primary mechanism for introducing elk to caves and rock shelter archaeological sites may have occurred at low frequency (27).
Elk are vagile mammals, with reported dispersal distances up to 600 km (51).Therefore, further range expansions of Rocky Mountain elk throughout the Sierra Nevada are not only possible, but contemporary, substantiated observations suggest it is likely.Although mean dispersal distances for Rocky Mountain elk are likely 42 km or lower (51), and the above longdistance dispersals are exceptional, they do seem to be occurring more frequently in the last decade.
In conclusion, although direct evidence suggests elk were never abundant in the Sierra Nevada historically, we find that the direct and indirect evidence, when taken together, indicates that elk likely inhabited at least the northern Sierra Nevada at least as far south as Lake Tahoe.
Long-distance dispersals to the central Sierra Nevada and recent colonization into its southernmost mountains, as well as a historical observer record of elk in the even more southern San Bernardino Mountains, raise the further question as to whether most of the Sierra Nevada is suitable elk habitat.Our findings should spur searches for additional physical evidence of elk in the Sierra Nevada during the historical period, utilizing novel approaches and technologies such as those that have recently expanded the recognized historical ranges of other species in California (16,52,53).Although we do not currently make recommendations for or against further elk reintroduction to the Sierra Nevada, it is clear that management activities that support and promote the recently established elk populations in Plumas and Sierra Counties may yield numerous ecological and social benefits.Sustained elk occupation in this wildfire-prone region may help in reduction of fire fuel loads (54,55), create new wildlife viewing and hunting (including indigenous hunting) opportunities (56), and reestablish an important prey base for apex predators and partial or obligate scavengers (57,58).Lastly, we recommend further research to better understand and manage the Sierra Nevada's (re)colonizing elk populations, including DNA analysis and application of satellite GPS collars to elucidate space use, habitat preferences, landscape connectivity, and identification of unoccupied habitats where range expansion by elk may naturally occur.
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Introduction
The California Department of Fish and Wildlife (CDFW) 2018 Elk Conservation and Management Plan depicted historical elk (Cervus canadensis) distribution as being absent from the Sierra Nevada of eastern California and western Nevada (Fig 1) (1).This estimate was based on twentieth century accounts written after possible extirpation of elk in the nineteenth century in the region (2,3).The nearest elk historically were thought to be Rocky Mountain elk (C.c. nelsoni) located in northeast California's southern Cascade Range and tule elk (C.c. nannodes) relegated to "the Central Valley and the grasslands and woodlands of central California's Coast

Fig 1 .
Fig 1.Estimated historical distribution of California's three elk subspecies (reproduced from Valley and Lake Davis, Plumas County, and Stampede Reservoir, Sierra County (CDFW internal data).Abundance and space use of these populations are not well understood, however elk occupation and reproduction in these regions indicate that habitat is suitable for elk, in at least the northern Sierra Nevada.Although the 2018 CDFW Elk Management Plan showed no elk in the southern Sierra Nevada, wildlife biologists since 2021 have observed elk, including three cows, with darker coloring consistent with Rocky Mountain elk, in the Sequoia National Forest/Lake Isabella area of Kern County at elevations above 800 m (A.Gwinn, CDFW and N.Kelly, USDA Forest Service, personal communication).These elk are thought to be dispersers from the Tehachapi herd, whose founders were Rocky Mountain elk translocated from Yellowstone National Park to the ranch of Rex Ellsworth next to Fort Tejon in Kern County's Tehachapi Mountains in 1967 (1).These conclusions are supported by three recent documented elk-vehicle collisions on Highway 58, near Tehachapi Pass, the boundary between the Tehachapi Mountains and the southern Sierra Nevada (A.Gwinn, CDFW and N. Kelly, USDA Forest Service, personal communication).There is no evidence that the tule elk removed from Yosemite Valley in 1933 and transported to the Owens Valley in Inyo County on the southeastern edge of the Sierra Nevada (46), have expanded their range westward, as is expected due to their high fidelity for more open range.
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