Introduction

The California Department of Fish and Wildlife (CDFW) 2018 Elk Conservation and Management Plan excluded the Sierra Nevada of eastern California and western Nevada from their map of historical elk (Cervus canadensis) range (Fig 1) [1]. This estimate persists in recent authoritative texts [2, 3] but was based on twentieth century accounts written after possible extirpation of elk in the region in the nineteenth century [4, 5]. 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 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 [5]. This strong habitat preference is likely related to physical and behavioral adaptations for drier, Mediterranean-climate conditions [6, 7]. 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 [4]. 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 [5, 8, 9]. Until then CDFW had disputed that this subspecies had existed in California, aligning with Murie’s perspective [4, 10]. However, McCullough’s findings were subsequently accepted by CDFW in their 2018 elk plan [1].

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Fig 1. Estimated historical distribution of California’s three elk subspecies.

Rocky Mountain elk (yellow) shown in northeastern California north of the Sierra Nevada, Roosevelt elk (blue) in northwestern California, and tule elk (pink) in California’s Central Valley and central coast regions. As shown, the Sierra Nevada were excluded from historical elk range. Reprinted from CDFW Elk Conservation and Management Plan under a CC BY license, with permission from CDFW, original copyright 2018.

https://doi.org/10.1371/journal.pone.0301665.g001

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 low wherever they occurred [11]. 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…” [11]. These two accounts, both in the Sierra Nevada region near the California and Nevada border, were based on an extensive search of historical newspaper and other observer records, but the specific citations were not listed. Nevertheless, the NDOW findings challenged prevailing thinking that elk never occupied the Sierra Nevada region.

Few contemporary studies are conducted with their primary goal the determination of the historical range of a given species [12]. 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 [13]. 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 [13–15]. These early venturers may have rapidly depleted elk in the Sierra Nevada before naturalists arrived to 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 [16]. 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 accepted 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 [13, 17] and the historical nativity of the California red-legged frog (Rana draytonii) to the Yosemite region [18]. 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) [19, 20], wolves have recently colonized both the northern and central Sierra Nevada, with packs currently (re-)established in Plumas and Tulare Counties, respectively–demonstrating that at least portions of the Sierra Nevada are suitable wolf habitat [21]. Similarly, a historical ecology study of the presence of wolverine (Gulo gulo) found historical observer records only in the southern subalpine and alpine Sierra Nevada [22], 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 [23].

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 [24], we evaluated multiple 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 [25]. 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.

Methods

Results

Historical newspaper accounts and other observer records

First, we located the two previously uncited historical newspaper references in the NDOW report, which described Sierra Nevada elk hunts in 1867 and 1868, respectively. The first historical newspaper account was printed in The Carson Daily Appeal in 1867 [35] 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 [33]. Zephyr Cove House was established by Andrew Gardenier, who homesteaded 65 hectares (160 acres) and built his inn in 1862 on Lake Tahoe’s southeastern shore to cater to silver miners. It was located at Zephyr Cove, Douglas County, Nevada [36]. The second elk hunt account was described in a pair of newspaper references in 1868 in the vicinity of Honey Lake Valley, Lassen County, California:

“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.

This account was printed in the Territorial Enterprise (Virginia City) in 1868 [37] and accompanied by a second report in the Virginia City Daily Trespass [38]: “a hunting excursion to Honey Lake Valley… to Doyle’s ranch, in Honey Lake Valley, which place they made their headquarters for a seven-days hunt… Hunting was done principally on horseback. For the first three days all the game killed was a noble elk, which weighed 404 pounds, having antlers at least four feet in length.”

- The Daily Trespass 1868.

Doyle’s Ranch still exists and is located in Milford, Lassen County, California on the southwest shore of Honey Lake, the terminus of the Susan River. Honey Lake Valley lies south of Honey Lake, 107 km north of Lake Tahoe. Both Honey Lake Valley and Lake Tahoe are south of the Susan River, placing both newspaper accounts in the Sierra Nevada (Fig 2) [39]. The two newspaper accounts are 122 km and 260 km south, respectively of CDFW recognized historical elk range (Modoc County’s southern border) [1].

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Fig 2. Map depicting locations of six nineteenth century elk observer records at four locations “H”: Honey Lake, Lake Tahoe, Round Valley, and the Bear River canyon in the Sierra Nevada.

Solid line depicts the boundaries of the Sierra Nevada in California and Nevada [39]. Two Late Holocene zooarchaeological records of elk “Z” are located in the extreme northwest Great Basin, just north of the Sierra Nevada. Habitat suitability is reflected by the contemporary presence “C” of breeding elk today in the northern Sierra Nevada south to Stampede Reservoir and the southern Sierra Nevada north to Lake Isabella. The dispersal of a bull elk to St. Mary’s Pass “C” south of Lake Tahoe is also noted. Images in map collected from public domain open sources (Esri, CGIAR, USGS).

https://doi.org/10.1371/journal.pone.0301665.g002

In addition to the two newspaper accounts of elk hunts mentioned in the 1998 NDOW Elk Management Plan at Zephyr Cove, Nevada and Milford (Honey Lake Valley), California, respectively, we located four additional historical elk observer records in the Sierra Nevada, bringing the total to six locations (Table 1).

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Table 1. Summary of historical observer records of elk hunts in the Sierra Nevada.

https://doi.org/10.1371/journal.pone.0301665.t001

The earliest record was in 1847 by Chester Ingersoll, who meticulously described his wagon train journey over the Sierra Nevada crest and down along the South Fork Yuba River then to Bear River Valley, Placer County, CA where he reported “Game plenty, bear, deer and elk…” [40]. Then, in 1874, two different elk hunts in Honey Lake Valley and Round Valley, Plumas County reported 15 and 13 elk taken, respectively [41, 42]. The last elk hunt newspaper record was described in two newspapers on Jan 8 and 9, 1875 where a dozen elk were taken in the vicinity of Honey Lake and described thusly: “Judging from the size of their bodies, they must have been monsters when living. In order to save a few dollars in freight the heads and legs were cut off of all the animals.” [43, 44].

After reviewing historical records for elk in the Sierra Nevada, we searched for other records in nearby regions that appeared reliable because they mentioned both deer and elk in the same observation. In the Great Basin adjacent to the Sierra Nevada’s eastern border, a report of elk in the Humboldt Sink was given by Thomas Jefferson Farnham, who explored California and Nevada 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.” [30]. Farnham also noted the occurrence of elk south of 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” [30]. 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 [45].

In contrast, Jedediah Smith, the first Anglo-American to cross the Sierra Nevada in 1827 at Ebbett’s Pass, 50 km south of Lake Tahoe, did not describe seeing elk [46]. Peter Skene Ogden, the discoverer of what is now the Humboldt River in 1828 and who explored it in 1829, also did not mention elk on his return from the Humboldt Sink over the southern Cascades to Oregon [47]. In the diary of Charles Preuss, who along with John Fremont were the first Anglo-Americans to see Lake Tahoe in early 1844 on Fremont’s second expedition, no elk observations were reported [33].

Discussion

Multiple direct and indirect data sources indicated that elk were historically present in the northern Sierra Nevada. Three of the six historical observer records establish the presence of elk with reasonable specificity, and two more of the six report elk in the same locations as the initial three suggesting they are bona fide. The 1867 Zephyr Cove, Lake Tahoe account of 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 [58, 59]. Although mule deer size is quite variable, the largest trophy animal taken in northeastern California had a live weight of 181 kg (400 lbs) [10]. The 1868 Honey Lake Valley elk hunt account described a “404 pound” (184 kg) elk but differentiated it from deer and pronghorn (Antilocapra americana). Further, the specific description of antlers “four feet in length” are unlikely to describe head ornaments for any ungulate other than elk [60]. Similarly, the 1847 Ingersoll description also differentiates deer and elk. Further, Ingersoll’s account provides detailed miles traveled from the Sierra Nevada crest, likely Donner’s Pass, through the South Fork Yuba River watershed then crossing to the Bear River valley en route to “Johnson’s”, which would have been Rancho Johnson, a Mexican land grant acquired by William Johnson in 1846 (now Wheatland, Yuba County, California). Ingersoll’s elk observation places the animals at about 850 m (2800 feet) elevation along the Bear River in Placer County, 1.6 km (1.0 miles) north of present-day Dutch Flat, California. This observer record is the furthest west in the Sierra Nevada, and at this elevation these likely were not the tule elk subspecies of the Central Valley [5]. The 1874 and 1875 accounts of elk killed at Honey Lake suggest that elk herd sizes may have been substantial, with 15 and 12 elk killed in each hunt, respectively [41, 43, 44]. These numbers were similar to the Round Valley 1874 elk hunt in Plumas County, where thirteen elk were taken [42]. It seems most likely that this Round Valley (there are several in the Sierra Nevada) is the former settlement established in 1863 in Plumas County, as the others are not populated places but rather just geographic features. Of note, the five historical newspaper accounts of elk hunts may have never been recorded if not for the discovery of silver ore in 1859 in Virginia City, Nevada. The prospect of wealth resulted in a population surge and the launching of several regional newspapers only 16 years before our records of elk cease in 1875. Similarly, the location of the 1847 wagon train observer record of elk might have remained inexplicable if not for Ingersoll’s punctilious diary entries [40].

The elk hunt historical observer records were supported by two Late Holocene zooarchaeological records in the northwest Great Basin, outside of but adjacent to the northeast Sierra Nevada, one in California and one in western Nevada. This physical evidence of elk in regions close to (just north of) the Sierra Nevada is supported by two reliable historical observer records also outside of but near the eastern and southern borders of the Sierra Nevada, respectively, by the early California explorer and naturalist, Thomas Farnham [30].

Indirect evidentiary records also support the historical presence of elk in the Sierra Nevada, similarly to the direct evidentiary records. 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 historical elk hunt there, may have an eponymous or other etymology unrelated to elk, the animal.

The paucity of historical observer records and Late Holocene zooarchaeological records are consistent with elk likely occupying the Sierra Nevada at 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.” [28]. The lack of museum records is likely related to the relatively late establishment of museum zoology collections in California, which opened 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 [13]. Another 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. Finally, Late Holocene zooarchaeological specimens may have been rare, because the region was sparsely populated by Native Americans, so the primary mechanism for introducing elk to caves and rock shelter archaeological sites may have occurred at low frequency [28]. These factors may have also led to a dearth of historical records of moose (Alces alces) in Nevada, another deer family member which has now colonized northeastern portions of the state with dispersals noted in the 1950s leading to an established permanent population over the last decade, although this hypothesis has not been verified [61].

Although we present evidence that suggests historical elk occupation of the Sierra Nevada, it is difficult to draw inferences regarding the ability of historical habitat conditions to support elk populations. However, it is clear that contemporary habitat conditions are suitable for supporting elk populations now and into the future. Habitat suitability in the northern Sierra Nevada is demonstrated by intergrade Roosevelt x Rocky Mountain elk herds exhibiting successful, sustained reproduction in the northern Sierra Nevada as far south as Stampede Reservoir [55]. However, because of their genetic admixture, these intergrade elk may be better adapted to higher elevations and novel terrain than their non-intergrade ancestors. This idea needs to be better explored through rigorous genetic analyses, such as next-generation sequencing, to be substantiated. Further northern range expansion of the nascent Lake Isabella herd in Kern County would suggest that the southern Sierra Nevada is suitable Rocky Mountain elk habitat. The recent recolonization of elk’s primary predator, gray wolves, in Tulare County, California in the southern Sierra Nevada, may also imply that elk are a missing component of the ecosystem [21].

Because elk are vagile mammals, with reported dispersal distances up to 600 km [62] further range expansions of elk throughout the Sierra Nevada are not only possible, but contemporary, substantiated observations suggest it is likely. Although dispersal distances for Rocky Mountain elk average 42 km or lower [62], range expansion to the central Sierra Nevada may occur in the near future as long-distance dispersals to the central Sierra Nevada have occurred with increasing frequency over 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 occupied the northern Sierra Nevada at least as far south as Lake Tahoe, as well as the northwest Great Basin, at least seasonally, in low densities, or both. Long-distance dispersals to the central Sierra Nevada and recent colonization into the southern Sierra Nevada, as well as a historical observer record of elk further south in the 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 [17, 63, 64]. 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 [65, 66], create new wildlife viewing and hunting (including indigenous hunting) opportunities [67], and reestablish an important prey base for apex predators and partial or obligate scavengers including California condors (Gymnogyps californianus) [68, 69]. However, given the juxtaposition of elk in the southern Sierra Nevada to the Owens Valley tule elk, there may be future genetic consequences and thus management implications to consider, particularly to preserve the genetic integrity of the endemic tule elk.

Human-elk conflict issues including fence damage and forage competition with livestock should be reported to the California Department of Fish and Wildlife’s Wildlife Incident Reporting (WIR) system and managed by established principles of adaptive management [70]. Consideration of wildlife overcrossings should be considered for Interstate 80 and other major highways, considering the proximity of the recolonizing elk in the northern Sierra Nevada. Lastly, we recommend further research to better understand and manage the Sierra Nevada’s (re)colonizing elk populations, including application of satellite GPS collars to elucidate space use, habitat preferences, landscape connectivity, identification of unoccupied habitats where range expansion by elk may naturally occur, and DNA analysis of untested herds such as in the southern Sierra Nevada.

Acknowledgments

The authors wish to acknowledge the thoughtful diligence of Robert “Bob” McQuivey, the retired NDOW investigator who in 1997 researched and discovered the historical newspaper and observer accounts of elk hunts in the Sierra Nevada, but whose detailed work remained unpublished for over a quarter century. We also wish to recognize Ian M. McGlory, MLIS, the Processing Archivist at Mathewson-IGT Knowledge Center, University of Nevada, Reno for his assistance locating images of historical newspaper accounts. We thank the CDFW biologists who provided the latest updates on the expanding elk populations in the northern and southern Sierra Nevada. This article was partially supported by CDFW, NDOW, and New Mexico Department of Game and Fish (NMDGF), with US Fish and Wildlife Service (USFWS), and are solely the responsibility of the authors and do not necessarily represent the official views of the CDFW, NDOW, NMDGF, or USFWS.