Great Herds: Elk

Roosevelt elk field sketch, Prairie Creek Redwoods State Park. Graphite pencil on paper.

On a sketching trip to the Pacific Northwest I stopped at Hansen National Wildlife Refuge along the Columbia River where a herd of Roosevelt elk mixes with White-tailed deer in lush meadows and wet Douglas fir forests. The month was October and the rut was still on. I sat by my spotting scope along a dirt road and watched a giant seven-point bull elk walk by, his antlers so long they nearly touched his rump as he threw his head back to bugle. Elk cow herds grazed nearby. As night fell, an eerie mist rose from the river, and pink clouds of sunset silhouetted the trees black.

Incredible sounds came from the forest that night: against a backdrop of hundreds of geese cackling in the distance, coyotes yelped, an owl hooted occasionally, and elk bugled--that indescribably wild whistle ending in a roar from the deep chest of an animal the size of a horse. And then came clashes of antlers, terrific cracking and echoing of two unseen opponents throwing their weight at each other in tests of strength.

This, I thought, must have been what primeval California was like when elk filled the forests, hills, and marshes. Now, what if I had been sitting on a hill overlooking a small cove in the past, in what would later become the city of San Francisco? My field notes, sketches, photographs, and memories may later combine into a reconstruction of a scene from Old California, as in the painting above. So often in developing a painting of the past, diverse material is needed from many geographical areas to put the picture together and fill in the missing pieces. Fortunately for elk in California, the picture is readily available and viewing elk in refuges and parks is relatively easy.

“Elk were so plentiful there that we made corrals out of elk horns picked up on the plains.”

--Jonathan Watson, market hunter in the 1860s in the San Joaquin Valley (Sleeper 1976).

Tule elk cow, Grizzly Island. Ballpoint pen on paper.

My original chapter title for the book, which was not used. Colored pencil on paper.

Coastal prairies would have had abundant tule elk sign: shed anlters, a few trails, and wallows where the elk took "dust baths." Detail of oil on panel, San Francicso hills view before European contact.
Herds of tule elk and pronghorn antelope in the Sacramento Valley before Europan contact. Valley oaks dot the grassland, and moist grassy-wetland edges provide succulent green vegetation for the herdfs. Oil on cotton rag paper.

Tule elk in the blue oak savanna of the South Coast Range. Oil on panel.

Before 1850 elk roamed huge expanses of grassland, oak woodland, and forest from Santa Barbara north into Oregon, from the coast over the Central Valley to the Sierra Foothills, and on into Northeastern California. No records exist for the Sierra Nevada, although elk may have wandered up the west slope in Summer to forage on lush meadows, and herds certainly wintered on the foothills during times when flooding formed extensive lakes in the Central Valley. The Los Angeles grasslands may have been too xeric to support elk, as no sight records or bones from archaeological middens are known for the region south of the Tehachapi Range.

Anchoring near the mouth of San Franciso Bay, “under a high and beautifully sloping hill, upon which herds of hundreds and hundreds of red deer, and the stag, with his high branching antlers, were bounding about....”

--Richard Dana, in San Francisco, December 27, 1835 (Barker 1994).

Elk range in the far west

Three subspecies of elk inhabited California, and through successful conservation efforts they still do. The endemic Tule elk (Cervus elaphus nannodes) is the arid-adapted elk of the oak-grass valleys and hills. It is lighter in color than other North American elk and has shorter antlers; old bulls sometimes develop palmate terminal points.

Tule elk have been labeled “dwarf elk” although this is misleading, and may be based on scientific descriptions of relict animals that survived the market hunting onslaught after the Gold Rush, or on elk translocated in 1933 to marginal habitat such as the Owens Valley, where the alkaline desert meadows are not prime elk habitat. Body size and antler growth are sensitive indicators of nutrition, and optimal development occurs when late Summer forage is of high quality (Geist 1991). McCullough (1969) tells of a Tule elk from Buttonwillow in the San Joaquin Valley transplanted to a Monterey golf course where it “grew to very large size” and looked more like a Rocky Mountain elk. H. C. Bailey, living in the Colusa area, described elk in the past as “magnificent,”sometimes having an antler spread of six feet across, and a dressed weight of 700 pounds (California Fish and Game 1931). In modern times, California Department of Fish and Game records show Tule elk bulls on Grizzly Island in Suisun Bay weighing up to 900 pounds. Cows weigh less, from 375 to 425 pounds (Suisun Ecological Workgroup 2003). I can only speculate that, in the past, Tule elk feeding all Summer in the lush freshwater marsh edges of the Central Valley could have grown to truly impressive proportions.

Roosevelt elk (C. e. roosevelti) are a large elk with darker pelage and heavy antlers that often have a crown of tines on the ends. Prehistorically they ranged from Sonoma County north into Oregon, inhabiting meadows and forest openings in the North Coast Range, to 7000 feet elevation (Thomas and Toweil 1982). They can still be viewed at places such as Prairie Creek Redwoods State Park.

The third subspecies of elk in California was the Rocky Mountain elk, a large-bodied type with long antlers that is famous today as the elk of Yellowstone National Park. It was noted in the Mt. Shasta region, along the Pit River, and in Siskiyou County, but was extirpated by 1873 (McCullough 1969). Since then, it has been reintroduced into Northeastern California (see chapter XX for more on this elk). This type has an average body weight of 755 pounds for bulls from the northern Rocky Mountains (Geist 1991).

Tuile elk

Endemic to California: Central California coast, Central Valley, Sierra Foothills, and South Coast Range to Santa Barbara

Roosevelt elk

Northwestern California into Oregon

rocky mountain elk

Northeastern California, Modoc Plateau, into the northern Great Basin and to Greater Yellowstone Ecosystem

Roosevelt elk, Prairie Creek Redwoods State Park. Felt-tip pen on paper.
Field notes of Roosevelt elk, while I was camping.
Roosevlet elk. Ballpoint pen on paper.

Tule elk were apparently abundant in Old California, and may have been the dominant big game animal in original grasslands ecosystems. Historical writings of trappers, hunters, explorers, and travelers frequently mention large herds.

Not all researchers, however, agree with the theory that pre-contact game numbers were high. Geographer William Preston, at California Polytechnic University, San Luis Obispo, represents a view that big game numbers, including elk, were prehistorically low due to overexploitation by native people. He explains that the large herds seen by almost all first contact travelers were “artificially high” because epidemics had swept through Native American populations even before Europeans came to California, epidemics released previously in Latin America by the first explorers from Europe. Preston argues that native people, as the “top predator” had overhunted big game animals before 1492, so that they would have been rare and marginal in early California, at least during the late Holocene. When native tribes became reduced by a “protohistoric plague” spreading from the south prior to missionization, game animal numbers expanded rapidly to “unnatural proportions” (Preston 1998).

Much of the evidence for this view comes from archaeological faunas interpreted in light of “Optimal Foraging Theory” and “Resource Intensification models.” Broughton (1999), for example, examined numbers of food animal remains in the Emeryville Shellmound in the San Francisco Bay region with the assumption that larger prey are more profitable for prehistoric people to hunt, allowing them a higher foraging efficiency. Bone counts for species of larger prey were said to decline over the period of use of the site, while smaller prey increased. This was interpreted as an indication of overhunting by native people, until such low numbers of big game were reached that a switch to more use of small game was necessitated.

Having been trained in the field of paleontology, I would hesitate to make so many assumptions and interpretations about a death assemblage of animal bones that may have had other factors acting on it--human behavioral and cultural traits affecting the choice of prey and selection of parts of a kill, unknown movements of new human cultures into the area, scavenging by other animals, changes in the biology of prehistoric animals, and largescale Holocene climatic fluctuations acting upon animal (and human) populations. In their work on environmental archaeology, Evans and O’Connor (1999) say, “Human food never representative of the full biological community....” This is a good caution to the heavy dependence on archaeofaunal remains for trying to recreate past ecosystems. Broughton admits that the number of elk bones in his samples were too small to show any significant trends in that species, and that the argument was based on deer numbers.

Other sources of evidence can also be equivocal. The debate about whether elk were rare or common prehistorically in the Yelowstone ecosystem still rages, with the same evidence resulting in opposite interpretations: Kay (1998) uses repeat photography to claim that no browslines on old pictures of conifers must indicate aboriginal overkill and rarity of elk; Meagher and Houston (1998), however, show old browselines in their repeat photography study, and give evidence that elk were not rare.

“In 1854 I found elk plentiful in the foothills west of the San Joaquin Valley, as well as in the tule swamp. Bob Dikemen and Lee Phillips were my hunting companions, and we practically finished up all the tule elk in that section between Martinez and San Joaquin City. The elk were originally ranging over the hills (along the east slope of the open foothills) as far south as Newman, going north as far as Grand Island. They were originally driven from the hills and valley land into the tules by the vaqueros rounding up wild horses and cattle, as well as by hunters. In 1854 they were nearly all driven to the tules, but the finding of horns of 6 to 8 prongs, all over the hills proved how plentiful they had been.

“I found no difference in size between these elk and the Oregon, Washington, Wyoming, and Colorado elk, and felt sure that the bulls would weigh 700 to 800 pounds. They struck me as weighing about as much as an average steer and their horns were fully as big as any elk I have ever killed or seen in other states.”

--H. C. Banta of Oakland, a market hunter from Missouri who located on the west bank of the San Joaquin River to hunt geese, ducks, and elk (Evermann 1915).

Field sketches, ballpoint and felt-tip pen on paper.
Anatomy study of an elk, field sketch, Yellowstone National Park. Pencil on paper.
Tule elk bulls lying around, San Luis Wildlife Area. Felt-tip pen on paper.

The assumption that epidemics reached California before European contact has also been disputed. Readings of many of the early Spanish explorers seem to depict a scene of abundant game alongside large native settlements; for example, Crespi recounts the reports of soldiers ahead of him telling of smokes in all directions around San Francisco Bay, and elk commonly seen; Fages in 1770 describes numerous villages in the Livermore Valley, a village of 600 people at Santa Barbara, and “entire herds of elk” in the Central Valley (Priestley 1937).

Each species responds in unique ways to hunting pressures, and this further complicates the picture. Elk, unlike some Ice Age big game animals such as giant ground sloths, had a long association with hunting societies, having originated in Siberia-Beringia in the Pleistocene and then spreading across Asia into Europe, as well as to North America tens of thousands of years ago. The elk of North America is one form of a huge cline of ecotypes that have adapted to a variety of habitats in the New World and Old, including those occupied by humans (Geist 1991). They may have coevolved with hunter-gatherers.

I am not saying Optimal Foraging Theory is wrong, and I do believe that some resource intensification did occur in the late Holocene. But in developing models of historical ecology, sources such as archaeological sites give only a partial aspect of the past ecosystem. As many other sources as possible should be integrated into the framework of data, such as discussed in Evans and O’Connor (1999). My own approach has been to follow the methods of lizard ecologist Walter Auffenberg, who in summing up his multi-volume studies on various monitor lizards of the world said, “Because the ecosystem is both the testing ground and the source of ecological principles, I have provided as much data as possible on both the environments of each of the species and the way in which each meets the challenges posed by its specific environment. In doing so I have deliberately provided more data than ordinarily might be the case in studies of this type. I am convinced that the development and testing of ecological generalities require an overflow of information on adaptive morphology, ecophysiology, systematics, and foraging and reproductive strategies” (Auffenberg 1994).

“Game was once very abundant-- bear in the hills, deer, antelope, and elk like cattle, in herds.... All are now exterminated, but we find their horns by the hundreds.”

--Brewer, camped in the Amador Valley near San Ramon, Alameda County in the 1860s (Brewer 1949).

Antler terms.

Past ecosystems are very difficult entities to imagine, as so much evidence has been lost or is open to wide interpretation. Even present, living ecosystems pose great challenges to decipher. Thirty years of intense research in the Serengeti Plains of east Africa, for example, have still not answered the many questions posed about its workings (Sinclair and Arcese 1995). What causes the complex wanderings of grazing animals, zebra and wildebeest, across the grasslands and savannas? Herding itself may be an antipredator strategy, but food seems to be the single most attractive pull to cause local movements and long-distance migrations. Grazers are constantly seeking forage with the highest nutritional value, the most succulent greenery.

For some time now I have thought about how Tule elk might have interacted with native grasslands in early California, what the seasonal movements of the herds may have been, how range ecology differed then from today. What follows are some of my speculations.

Grassland slope and aspect phenology drawing, from years of field observations and notes, Tilden Regional Park, Contra Costa County, California. Ballpoint pen and colored pencil on paper. Tule elk may have taken advantage of the greening-up of vegetation on different slopes and aspects during the rainfall year.

What is slope?

Slope refers to the extent that a soil surface has an incline relative to the horizontal. In percentage terms, slope represents the elevation that occurs between two different points. When the terrain is flat, there is no slope or aspect.

What is aspect?

Aspect refers to the compass direction a land surface faces. For example, a northern aspect slope faces more northwards, receiving less sunlight than southern aspect slopes. This greatly influences microclimates, moisture content of soils, and vegetation types, as well as seasonal green-up. In vegetation community science, north-facing slopes and south-facing slopes are very imporatnt to observe.

Slope-aspect map

Take a look at this informative slope-aspect map by GIS geography at

Elk would have taken every opportunity to follow the green-up on certain cool moist slopes at the end of summer, and stay warm on south-facing slopes during cold winter periods. Before the Age of Fences.

Chart showing the green periods of native bunchgrasses, perennial creeping wildrye (Elymus triticoides), and wildflowers/forbs in the East Bay Hills. Summary after years of collecting observations and making notes.

A wildlife photographer friend of mine traveled to Africa most years for two or three months, and brang back stories of sitting by a waterhole to wait for hours for antelope or elephant to come to drink, tripod and camera at the ready. I listen fascinated, longing to go myself to witness great herds of wildebeest, or hear lions roar at night. But then my mind turns to California: I imagine myself taking a sketchbook and a backpack, and hiking over golden grassy hills say 400 years ago. I would have had quite a safari looking out for grizzlies, listening to wolves howl at night, and seeing large herds of elk come along a trail to drink at a marsh pool.

Reconstructing the food habits and movements of prehistoric elk is only guesswork, but clues can be found by studying elk today, as well as the relict native plant communities that survive.

Elk feed predominantly on grasses and sedges, with forbs very important in the spring and summer, and the leaves of shrubs and trees also favored throughout the year. In other words they are successful generalists, taking the best plant food available. Tule elk skulls, although shorter than those of other elk, have longer toothrows of grinding molars and premolars for chewing the fibrous plants of the semiarid California prairies (McCullough 1969).

California had a mosaic of patches of bunchgrasses, rhizomatous grass swards, and some annual grasses. Elk moved through this mosaic choosing the most palatable species with the newest growth. Four days after the first rains in October, Purple needlegrass (Stipa pulchra) begins to green up, starting perhaps on ridgetops and warm south- and west-facing slopes. Elk may have concentrated in these spots to dine on green blades. During the winter rainy season, green bunchgrasses would be widespread, and elk could have dispersed over hills and plains. They may have selected particularly tender species ahead of coarse grasses: the small tufts of bluegrass (Poa secunda) that once were abundant on slopes and flats of the California prairie; the soft fine blades of Junegrass (Koeleria macrantha), and the moist bunches of Bromegrass (Bromus marginatus). The last grass, when grazed down early will produce a luxuriant aftermath of foliage that is relished by grazers even in the dry season.

Tule elk bull dining on the new green grass grouwth of purple needlegrass bunches, a month after a fire. Elk will travel long distances to forage on these succulent "ice cream" plants, and before the era of livestock fencing elk herds probably wandered across the valleys and hills of central California in search of new burns. Indigenous people therefore used cultural burns (#GoodFire) to attract elk for hunting, and manage elk habitat. This was disrupted in the early 1900s when colonizing federal agencies disrupted this ancient pattern of grassland fire management to aid the elk. Oil on cotton rag paper.

“The coast we were passing is formed of mountains of moderate height, covered with grass, at that time somewhat parched; in the ravines we saw clumps of oaks. From time to time we descried large deer [elk] herds. They were wandering in bands over these sloping pasture grounds....”

--Duhaut-Cilly, 1830s, sailing in San Fransisco Bay from Angel Island to San Rafael (McCullough


Herd of tule elk roaming across a summer blue oak ssavanna, in the South Coast Range. This would have been a common sight in Old California. Oil on panel. This painting was based on my field observations of tule elk near Sunol Regional Wilderness.

During the spring elk probably turned to feeding on the abundant wildflowers that cloaked the prairies. In elk forage studies, I find long lists of California native annual and perennial forbs eaten by elk, from buttercups (Ranunculus californicus), California poppy (Eschscholzia californica), owl’s clover (Orthocarpuss pp.), Soap plant (Chlorogalum pomeridianum), to lupine (Lupinus spp.), brodiaea (Brodiaea and Dichelostemma spp.), and many others.

A feast of high-protein, nutrient-laden forbs allowed elk to put on weight before the summer dry season, and shifted the grazing pressure away from the bunchgrasses as they put out seed stalks.

When the grassy hills dried out and spring wildflowers turned to dust, elk shifted again to feed on green willow leaves in valleys, oak leaves, and leaves of small trees andshrubs such as blueblossom (Ceanothus spp.), currant and gooseberry (Ribes spp.), wild rose (Rosa spp.), and elderberry (Sambucus spp.). Summer-green wildflowers may also have been important, such as the multitudes of aromatic tarweeds (Holocarpha virgata, Madia spp., Hemizonia spp.) that grew among drying bunchgrasses, the buckwheats (Eriogonum spp.), and asters such as the purple-flowered Aster chilensis. Purple needlegrass when dry cures well in the summer, and may have been grazed by elk as a source of carbohydrate energy--good quality hay.

Local movements of elk down to valley bottoms may have occurred in the dry season as extensive patches of Creeping wildrye (Elymus triticoides) would have provided much green forage. Marsh edges around San Francisco Bay, and especially in the Central Valley, would have provided an abundance of luxuriant vegetation, green water-edge grasses such as Spike bentgrass (Agrostis exarata), Sloughgrass (Beckmannia syzigachne), witchgrasses (Eragrostis spp.), Ditchgrass (Papsalum distichum), and Reedgrass (Phalaris arundinacea), as well as sedges, rushes, green forbs, and willows.

In the fall, elk could gorge on acorns back in the uplands, and travel to early green-up areas as the rains began, to start the cycle over. Indigenous land managers would have added a crucial food resource: burned fields. Many tribes lit fires in grassland and chaparral areas, as discussed in the Fire Chapter. Fire removed old dead leaves on perennial bunchgrasses and stimulated new growth of green leaves, even in an otherwise dry summer landscape. These green patches would have been highly attractive to elk, deer, and antelope, producing high quality forage during a time of more limited nutritional resources. A mosaic of burned and unburned grassland would keep elk moving.

Unlike many modern cattle operations where stock are fenced into small areas, and not allowed to wander to seek the best forage patches, and where the wandering instinct has even been bred out in favor of meaty animals, elk in early California probably seldom grazed the same area more than once or a few times a year. A complex deferred rotation grazing system may have been in effect, allowing individual plants a chance to recover (Houston 1982).

Field sketch at Sunol Regional Wilderness of cattle walking into a herd of tule elk who werer lying in their beds--the elk jumoped uop and walked away. Here the cattle behaviorally impacted the native tule elk, and disrupted their use of the land. Ballp[oint pen on paper.

Even so, the reports of huge elk herds, sometimes numbering in the thousands, must have left some imprint on the land. If I could take a walk over the hills in Old California, I would have seen conspicuous trails leading to watering areas, some trampling and heavy grazing in favored spots, browselines on the willow thickets in places, and large wallows where bull elk during the rut would roll and paw the mud. Purple needlegrass on heavily used areas like ridgetops might have formed a zootic climax, responding to elk grazing by reducing the diameter of individual bunches.

An indirect indication of moderate grazing pressure by elk on California’s native grasslands comes from one of the earliest sightings of Tule elk, by Sir Francis Drake as he landed somewhere in the Point Reyes area. His sailors on making shore found large herds of "very large and fat Deer," which newcomers would have recognized as similar to their Red deer of Eurasia, basically the same species as North American elk. These would not have been the much smaller black-tailed deer. Drake's landing somewhere on the central California coast around what is now Point Reyes National Seashore, allowed his crew to witness large herds of tule elk, an amazing snapshot of life in the summer of the 1500s:

“Our necessarie business being ended, our Generall with his companie trauailed vp into the Countrey to their villages, where wee found heardes of Deere by 1000. in a companie, being most large, and fat of bodie.

“We found the whole Countrey to ba a warren of a strange kind of Connies, their bodies in bignes as be the Barbarie Connies, their heads as the heads of ours, the feete of a Want, and the taile of a Rat being of great length: vnder her chinne on either side a bagge, into the which she gathereth her meate, when she hath filled her bellie abroad. The people eate their bodies, and make great accompt of their skinnes, for their Kings coate was made of them.”

----Richard Hakluyt, of Sir Francis Drake’s voyage, in “The Principal Navigations, Voiages and Discoveries of the English Nation...” 1589 (Orr 1950).

The "Connies" were probably our California ground squirrels (Spermophilus beecheyi), which have been eliminated from the Point Reyes coastal area and many other Bay Area grasslands by pesticides and a war on native rodents in favor of commercial livestock grazing.

California ground squirrel sketch. Ballpoint pen on paper.

As the rut begins in July, an exciting event to observe today, the big bulls develop long black neck hair, contrasting with their light body fur. They seek out cow herds to join, chasing away lesser males. The organ-like bugle is now heard often, its long ending squeal carrying far over the grasslands.

the rut

Field sketches at Grizzly Island. Ballpoint pen on paper.

In August and September the rut peaks, and master bulls can be seen charging challengers with neck extended and head tilted up, showing off mighty racks of antlers. If the intruder does not run off, the two may walk parallel to each other in display, until one suddenly wheels to face his opponent and they smash their antlers together with terrific force.

The rut, field sketches at Tomales Elk Reserve in Point Reyes Natiuonal Seashore. Ballpoint pen on paper.

By October the rut is winding down. Harem bulls are exhausted from battling rivals, cows now begin to gather into larger winter herds. Bulls group back into bachelor herds again.

Elk bull herd at Tupman Elk State Reserve in the San Joaquin Valley. Ballpoint pen on paper.

“It was now fifteen miles to the San Joaquin River, and a level plain lay before us. When our road turned into the river bottom, we found the water too deep to get through safely, so we concluded to go on and try to find some place where we could cross. On our way droves of antelopes could be seen frolicking over the broad plains, while in the distance were herds of elk winding their way from the mountains towards the river for water. When far away, their horns were the first things visible, and they much resembled the dry tops of dead pine trees, but a nearer view showed them to us as the proud monarchs of the plain.”

--William Lewis Manly, heading across the San Joaquin Valley in Spring, 1850 (Manly 2001).

Tupman Elk State Reserve.

When watching big game one May in Yellowstone National Park, a lone elk cow passed below the hill I was sitting on. With binoculars up, I sketched her (above), her mouth open as if nervous. She looked about constantly until she entered a grove of Douglas fir and brush in a ravine--she appeared to be seeking a hidden spot in order to drop her calf.

I wondered at how difficult this must be, with grizzlies and a wolf pack in the vicinity, both sniffing the ground for every sign of calving. Such scenes must have happened many times in Old California.

The elk mother stays alone with her new spotted calf, away from the herd, for two to three weeks. She feeds as it lies hidden in brush or tall grass. When all is safe, the cow gives a special call and the calf jumps up and runs over to nurse. Then it seeks cover again.

By the calf’s second week it starts to feed on green grass, and soon it is strong enough to run. The mother and young then join the herd. The calf looses its spotted coat by July or August and is weaned by September or October.


Days-old elk calf hiding in sagebrush while the mother grazes nearby, ever vigilant. Yellowstone National Park, Wyoming. Ballpoint pen on paper.

Cautious mother Rocky Mountain elk, Yellowstone National Park.
A passing grizzly does not scent the newborn elk, and moves by as elk cows watch. Yellowstone National park. Felt-tip pen on paper.
A Rocky Mountain elk cow and calf exit the scene. Yellowstone National Park. Felt-tip pen on paper.

The cycle begins again as the bull elk gradually regrow new antlers, nourished by "velvet" skin through the summer.

Tule elk bulls growing new antlers, covered in velevt. Tomales Elk Reserve. Ballpoint pen on paper.
Tule elk near Sunol Regional Wilderness. Ballpoint pen on paper.

“ herd of Elk had a grand appearance containing more than 2000 Two thousand head and covering the plain for more than a mile in length....”

--James Clyman in 1845 on the west side of the San Joaquin Valley near Livermore (McCullough 1969).

elk diets

Field sketches at Sunol. Ballpoint pen on paper.

Many native plants have adaptations for seed dispersal by hitching a ride on the fur of passing elk or deer. Most tarweeds have achenes with gland-tipped hairs to stick to fur (and clothes). Cocklebur (Xanthium strumarium) has hooked spines on its seeds, and may have caught on elk fur around waterholes. The native Rancheria clover (Trifolium albopurpureum) has numerous hairs on its seeds to catch on fur. Many native grasses also are easily caught in fur from long bristly awns on their seeds: needlegrasses (Stipa spp.), Blue wildrye (Elymus glaucus), and Meadow barley (Hordeum brachyantherum).

In the search for feeding grounds, the question of long-distance migration has come up. Many populations of Rocky Mountain elk are migratory between high mountain ranges in summer, and lower valleys in Winter when snow accumulates. Eastern elk were apparently non-migratory (Geist, in Thomas and Toweill 1982). Roosevelt elk in the Siskiyou Mountains, according to settlers, migrated to winter ranges at lower elevations (Adams, in Thomas and Toweill 1982). A tally of dates and locations of Tule elk sightings by early travelers shows no pattern of migration, however. Elk were seen on coastal hills in July as well as in December. Elk were found abundantly in the freshwater marshes of the Central Valley in April, June, and October. Similarly in the Coast Ranges, they seemed to occur at all seasons. Thus elk in much of California probably moved about locally in response to rains, burning, wildflower blooms, and wetland green-up, instead of making long migrations like Serengeti wildebeest.

Elk bull in velvet, Tomales Elk Reserve. Ballpoint pen on paper.

In my wanderings through the oak and grass canyons of Tilden Regional Park near Berkeley, I found an odd land feature. It was a depression about 30 or 40 feet long, as if scooped out by a bulldozer. But it was old, with mature Creeping wildrye growing over its banks, and small springs seeping water near it. A creek ran next to it, but this was no erosional feature. After looking at it for long periods, and noticing whitish soil in it, and even a strange species of tarweed growing at the spot that I had seen nowhere else, I decided that this could be an old mineral lick. Perhaps deer and elk over the centuries pawed it apart to supplement their diet with calcium, magnesium, or sodium, minerals rising out of the ground in solution from a fault. These are ideas I speculate about while hiking.

And so life went for the elk herd in Old California, actively feeding before first light. Each individual lay down to ruminate-- chew their cuds for two hours or so. Then more feeding. At midday they idled about their bedding area, and in the afternoon feeding resumed, continuing intermittently through the night. Feeding and resting occupied 90% of the elk’s day.

The seaonal cycle of elk in California can be summarized in a general way in the chart below.

The elk year

“We have been a month here and we could not have fallen on a better place to pass a part of the dead winter season when nothing could be done in the way of trapping on account of the height of the waters. There was excellent feeding for the horses, and abundance of Animals for the people to subsist on, 395 elk, 148 deer, 17 bears, & 8 antelopes have been killed in a month which is certainly a great many more than was required, but when the most of the people have ammunition and see animals they must needs fire upon them let them be wanted or not. The Animals for a considerable time back have been in general very lean, indeed they could not be expected to be otherwise being hunted without intermission.”

--John Work, leading a Hudson Bay Company expedition to the Marysville Buttes area in 1833 (McCullough 1969).

Loss and recovery

This daily and yearly cycle that went on in California for tens of thousands of years was broken, however. In the late 1700s and into the 1800s ranch eros in Spanish California hunted elk on horseback, lassoing them like cattle and taking their hides and rending their fat. Elk numbers did not seem to decline and elk may have been able to hold their own in this period, coexisting with cattle and horse herds on open ranges.

Fur trappers took elk also, from the 1820s to 1840s, apparently resulting in decreases in parts of the Sacramento Valley (McCullough 1969).

Not until the market hunting period during and after the Gold Rush did the elk population crash. Intensive hunting to supply meat to California’s booming population led to the disappearance of elk from the San Francisco Bay area by the 1860s. Roosevelt elk were driven from Sonoma County by about 1870 (McCullough 1969, Skinner 1962). The last stronghold of Tule elk was in the vast marshes of Tulare and Buena Vista Lakes in the San Joaquin Valley, and they probably would have gone extinct there were it not for a rancher, Henry Miller, who in about 1874 found a few remaining elk on his large cattle ranch when draining a marsh--the last Tule elk in the state. He decided to protect them, and this initiated recovery efforts by the California Department of Fish and Wildlife (Evermann 1915, McCullough 1969).

Elk were transplanted gradually to preserves, parks, and ranches across the state, and their numbers grew so that today there are now around 3000-- a far cry from McCullough’s (1969) estimated 500,000 before 1800. Tule elk, despite their population growth, still carry the legacy of this bottleneck in their genes. The diversity of genetic types has been lost, and all Tule elk today show their relatedness directly back to Henry Miller’s ranch individuals (Matocq et al. 2002). I can only speculate how elk populations across California may have differed slightly from one another. Subfossil bones from Pacifica on the coast show one skull character that approaches Roosevelt elk, yet other characters are like Tule elk from the San Joaquin Valley (McCullough 1965). We have probably lost an enormous amount of diversity in California elk.

Two elk bulls with growing antlers in velvet, grazing on a meadow with a mule deer. Yellowstone National Park, spring. Felt-tip pen on paper.

But overall the elk story in California is a cautiously successful one. Biologists, conservation groups, hunters supportive of conservation measures, parks, and private ranchers and land owners who do not mind elk herds wandering on their land, have all contributed enormously to the growth of elk numbers. Yet the genetic bottleneck continues and poaching on private lands is an ongoing problem.

Climate change has led to extreme droughts that have caused die-offs in tule elk trapped behind fences at the Tomales Elk Reserve in Point Reyes National Seashore, most recently in the very dry summer of 2021. See more at Save Point Reyes, Harvard Law Clinic (which sued over the drought kill), and Mission Rewild.

“At times we saw bands of elk, deer and antelope in such numbers that they actually darkened the plains for miles, and looked in the distance like great herds of cattle.”

--Edward Bosqui, between Merced and Stockton in the San Joaquin Valley (McCullough 1969).

George C. Yount exclaimed about California: “The deer, antelope and noble elk...were numerous beyond all parallel.”

--Quoted in Charles L. Camp, The Chronicles of George C. Yount,” 1923 (Dasmann 1965).

“I have no where seen game as plentifully as in this valley. We killed an antelope in the morning. We could frequently see herds of deer and elk in different directions around us, as well as wild horses.”

--A. B. Clarke visiting the Central Valley in 1852 (Dasmann 1965).

Elk viewing is easily done: Point Reyes National Seashore, Grizzly Island Wildlife Area, San Luis National Wildlife Refuge, Cache Creek Wildlife Area, and Praire Creek Redwoods State Park and Redwood National Park (Roosevelt elk), among others. In these places one can feel the heat waves in summer bounce off the backs of elk, watch a kingbird flycatch among their hooves, follow the tracks of a herd through the mud of a winter valley, and hear the wild bugle of an unseen bull echo through the oaks at dawn.

Bison in California

Rocky Mountain elk and bison field sketch, Yellowstone National Park. Ballpoint pen on paper.

Besides elk, California had other big “hoofstock ”: the historic Bison of the Modoc County region. On the northeastern sagebrush-grass plains and juniper hills the only certain population of Buffalo (Bison bison) in California once roamed. Herds probably mixed with Rocky Mountain elk, Pronghorn antelope, and Rocky mountain mule deer, as well as White-tailed deer. But an important question always dogs me while walking around California grasslands: Why were Bison found only in this corner of the state during the last several thousand years, and not all over?

First let us look at the historic northeastern buffalo herds. I have spent a lot of time camping and hiking the Modoc Plateau (Modoc and Lassen Counties) because of its remant antelope herds and vast marshes filled with waterfowl, and I have come to know the peculiar type of cold-arid grassland that exists here. Mixed with pungent Big sagebrush (Artemisia tridentata) and scattered dark green Western juniper (Juniperus occidentalis) is a grassland of Bluebunch wheatgrass (Pseudoroegnaria spicata), Bluegrass (Poa secunda), Idaho fescue (Fesctua idahoensis), Praire junegrass (Koeleria macrantha), and Thurber needlegrass (Stipa thurberiana) among others. This Great Basin community actually stretches in varying form in a broad arc across the interior West to the Rocky Mountain region, where it can be found in places like Yellowstone National Park (see map). Bison are grazers and would have selected for these grasses over shrub and tree browse. A good place to see this habitat in fair condition is at Lava Beds National Monument in Modoc County.

Map showing ther main Holocene bison range on the yellow plains grassland on the right. Potential secondary ranges of bison in smaller more scattered herds are shown west of the Rocky Mountain axis in intermontane habitats.

Bison bull, yellowstone National Park. Ballpoint pen on paper.

Bison bull studies in the field, Yellowstone National park. Ballpoint pen on paper.

Bison herds were small and apparently transitory here, as in much of the West outside of their Great Plains core range. What kind of bison these were is a matter of debate. Although little evidence is left after colonial expansion and over-hunting, the far-western bison may have been a different subspecies than the plains bison (B. b. bison), called the mountain bison or wood bison (B. b. athabascae). It ranged in Oregon, Washington, Idaho, northern Nevada and Utah, and up into Canada, and was noted by early Euro-American travelers as darker in color and somewhat larger in size, living in small bands of 5 to 30 animals in mountainous areas, timbered lands, meadows, and valley prairies (Meagher 1978, Soper 1941). Today this bison can only be seen at Wood Buffalo National Park in Alberta. Jerry MacDonald, however, after studying large series of bison skulls across the continent decided that skull measurements from Malheur Lake, Oregon were closer to the plains bison (B. b. bison) than the wood bison (MacDonald 1981). This would make sense judging from the map of the Bluebunch wheatgrass steppe reaching from Wyoming and Colorado across to northeastern California.

The naturalist C. Hart Merriam spoke to native people in northeastern California who said their fathers had killed these animals, and told about how the bison came in small bands from the north, perhaps seasonally (Merriam 1926). Paiutes remembered bison once living around the lakes and marshes of Malheur Lake, a habitat similar to parts of the Klamath Marshes (Allen 1942). Bison were found around Great Salt Lake in Utah, and a bison skull was found in Iron County in far southwestern Utah by Cedar City. One-hundred twenty-eight miles west of this bison bones were found in an Indian rock shelter in Lincoln County, southeastern Nevada (Bradley and Brechbill 1966). In late prehistoric times bison were seemingly pushing the edges of the driest desert belts.

In northeastern California a hunter in 1871 told of finding weathered buffalo skulls as far west as the Sierra Nevada (Merriam 1926). The Pit River tribes and Northern Paiute knew the bison well, telling how they used to hunt them with bows and arrows, although they said it was a difficult feat (Riddell 1952). Bison were found in Alturas Valley, Surprise Valley, the Madeline Plains, Hot Springs, Horse Lake Valley, Eagle Lake, Honey Lake, and a small permanent herd at Pine Creek Valley (Merriam 1926). The Smoke Creek Desert in adjacent northwestern Nevada has many place names such as “Buffalo Meadows” and “Buffalo Spring.”

Bison were extirpated in Lassen County about 1830 as settlers invaded the area (Riddell 1952). A buffalo wallow was described on the west shore of Honey Lake, with a trail leading southwest towards Plumas County over a low pass (about 5,000 feet in elevation). Bison were said to have taken the trail over this part of the Sierras, but no one knew where it led (ibid.).

Did bison cross this low pass into the western California Purple needlegrass habitats? Most biologists writing of California large mammal distribution deny that bison were found in California west of the Cascade-Sierra divide during the Holocene. But I have kept an open mind. Hiking the gentle Plumas County Sierras where the transition begins with the Cascade volcanoes to the north, I found open Ponderosa pine forest with a good grassy understory -- not a big barrier to bison considering that in other parts of the continent they regularly travel through conifer forest, as in Yellowstone National Park and in Canada.

Although I am not a specialist in the exact Spanish translations and meanings of the words used by early California explorers, there are some curious references to animals that may not be simply elk: in the 1770s Pedro Fages, traveling from the San Fransisco Bay to the Central Valley looked upon the Sierras and told of reports of “numerous wild buffaloes living in the dense parts of the forest” there (Priestly 1936). Father Crespi, too, on October 11, 1769 near the Pajaro River (Santa Cruz-Monterey Counties) reports that soldiers showed him:

“various tracks that looked like cattle, and we suppose they are buffalo. I did not see any of the beasts myself, but the scouts reported that along the shore hereabouts they saw as many as twenty-one beasts together, of all colors, with calves at their feet like cows; also, that they had seen deer or stags, very big-bodies and with very large spreading antlers...” (Paddison 1999).

What were these beasts, apparently not elk as the soldiers identified those separately? Were they wild cattle already spread from colonies in the Southwest? Or were these a small herd of wood bison, very early hunted out of western California?

Perhaps western California grasslands were too arid for bison, which are found today mostly in areas with higher summer rainfall. But bison have in near-recent times trickled into arid grasslands in the American Southwest. Historians found scattered accounts of bison or bison parts such as hides and horns mentioned in northern Mexico before 1600 (Reed 1952). A document dated to 1806 by an official at Monclova, Coahuila tried to ban settlers from their incessant hunting of bison, called “cibolo” (Dobie 1953). There were also bison reports from Chihuahua (ibid.) in semidesert grassland with cabbage palm (Sabal) and mesquite (Prosopis). Bison skeletal remains were found in various archaelogical sites west of the Rio Grande before 1200, giving evidence to me of a range contraction perhaps related to the climatic fluctuations of the Medieval Warm Period and Little Ice Age -- during times of increased moisture the bison bands moved beyond the fringes of their core range as grassy areas expanded, but during dryer phases shrunk back. A dramatic increase in the number of bison bones recovered from archaelogical sites corresponds with greater grass abundance in the Great Basin sagebrush steppe during the cooler moister period from 1,500 to 1,200 BP (Kinney 1996). California bison could have been irregular wanderers, as many game herds probably were before fences and park boundaries. Father Francisco Garces travelling up the Colorado River into Arizona in 1775 to 1776 reported that in Yavapai country by Cataract Creek and the Grand Canyon in Coconino County the Indians had killed a buffalo, as well as stray cattle and deer, and shared the meat with him (Galvin 1967).

Another possibility is the long-term multi-millenial expansion that bison may have been undergoing, well-described by paleontologist Jerry MacDonald (1981). Bison originated in Eurasia and during the late Pleistocene traveled across Beringia to North America where they flourished as several species of Ice Age types, including Bison antiquus which was abundant in California. Bison did not all go extinct with the rest of the megafauna but certain populations did show range shrinkage and skeletal abnormalities due to inbreeding in small isolated populations during the period from 12,000 to 6,000 years ago -- a bottleneck from which they survived. During this time bison body size decreased and horn size became smaller. Careful dating of skeletal remains through a series of maps over the last 12,000 years shows the range of bison shrinking from continental proportions to the northen Great Plains by 7,000 to 5,000 years ago -- their core area of survival during the Xerothermic mid-Holocene climatic pressure. After 5,000 years they may have undergone a slow range expansion as the climate became generally moister and cooler, and this is when our modern Bison bison appeared. MacDonald says their southward expansion accelerated by 3,000 years ago, and by 2,500 years ago they began to colonize outlying areas in a great range enlargement, to areas such as the Great Basin steppes. The ancient bison had successfully adapted and evolved to the radically changing post-Ice Age climates and plant communities.

San Francisco Bay 40,000 years ago during the late Pleistocene glacial phase, as I recontruct it based on research, field work, and imagination. At this time, the ocean was far out on the continental shelf due to continental glaciers holding massive amounts of water; therefore the bay was a valley, where the San Joaquin-Sacrmaneto River flowed through and out the Golden Gate, seen in the distance beyond a forested hill which is now Alcatraz Island. Hereds of fossil bison and horses graze here: Ice Age bison (Bison antiquus)--which evolved into our modern Bison bison--and the extinct Western horse (Equus occidentalis), which might have had characters of the partially-striped quagga (an extinct equid of Holocene southern Africa, named after its loud vocal call). Native bunchgrasses and sedge patches are shown with coast live oaks and bays, and Douglas fir forests. San Francisco would be now on the distant ridge on the left. 1991, acrylic on paper, 19.5 x 25.75 inches.

Bison populations over the last few thousand years apparently fluctuated with long-term droughts and cold storms but they were filtering in small numbers east of the Mississippi River and into Mexico, northern Arizona, the grassier parts of the Great Basin, and possibly even western California. Evidence indicates that this may have continued steadily were it not for Euroamerican settlers and hunters who cut this great expansion short. Bison in eastern North America were apparently in the process of range expansion when they were first encountered by Europeans. They may not have grazed east of the Mississippi until A.D. 1000 (Shay 1978). I believe that bison may have been irregularly wandering into northeastern California (and perhaps father west) in small groups for millenia.

Although I disagree with them, some biologists offer different scenarios -- Valerius Geist for one (Geist 1996) theorized that the waves of European epidemics that reduced the Indian population in the 1600s freed up the bison and other big game from heavy hunting pressure to greatly expand in number, and that the early reports of great herds in a vast wilderness of America were unusual; they argued that for most of the Holocene Native hunters would have kept bison at a low population level. I am of the opinion that bison were much more difficult to hunt than this theory proposes (especially without horses), that overhunting of big game did not happen until the introduction of guns and a worldview of resource-wasting, and that the influence of climate and evolutionary events were the deciding factors.

But then the bison were gone, hunted out by the colonizers. Millions of buffalo were killed by Amercian settlers and travelers between the 1860s and 1880s in their core range, creating a second Holocene bottleneck according to MacDonald, which probably resulted in the loss of certain phenotypes (physical types) and behaviors. Their range has been fragmented and artificially regulated ever since. But they still survive.

Field sketch, bison with calves playing, and Rocky Mountain elk. Yellowstone National park. Ballpoint pen on paper.
Field sketches of bison bull at Yellowstone National Park. Ballpoint pen on paper.

I pondered this while bison-watching in Yellowstone National Park. Small groups of young bulls scattered among the cottonwoods with herds of elk on the flat. A group of cow bison with many orange calves grazed nearby. Two young bison romped together in play, loping after the elk and scattering them. Then the two butted heads. In my mind I could translate this scene into Modoc County 500 years ago, to places such as Honey Lake, Alturas, or Lava Beds where I had spent so many hours watching and sketching Mule deer and antelope.

Whether bison would still be living wild in the Modoc plains and lava rimrock lands before the age of the rifle is now an undecipherable puzzle. Were they expanding their range naturally into California? Would climate change have affected them, or increasing drought pushed them back out? Historical accidents such as hunting out elk or bison from a region, a disease hitting a species, or a sudden climatic turn, can ripple across a landscape and change the interrelations of several other species. Our next chapter on predators of Old California shows this in particular.