Journal of Mammalogy, 90(1):74–92, 2009
NOBLE MARTEN (MARTES AMERICANA NOBILIS)
REVISITED: ITS ADAPTATION AND EXTINCTION
SUSAN S. HUGHES*
Northwest Archaeological Associates, Inc., 5418 20th Avenue NW, Suite 200, Seattle, WA 98107, USA
The cause of extinction of the noble marten (Martes americana nobilis), as well as its taxonomic position, has
been the subject of debate in recent years. This extinct marten, a close relative of the extant American marten
(Martes americana), is known from 18 sites in western North America, most dating to the late Pleistocene.
Because boreal fauna were associated with the late-Pleistocene noble marten, researchers generally believed that
it inhabited boreal forests like the American marten, and competition between the 2 may have caused its
extinction. Recent discoveries of noble martens associated with xeric fauna from Holocene contexts have called
these assumptions into question. I explore the adaptation and habitat of the noble marten with an analysis of its
faunal associations and find-site locations. The analysis suggests that the noble marten was adapted to open,
mesic grasslands in montane foothills, and was likely not sympatric with the American marten. I also introduce
a new Holocene noble marten specimen, a right mandible dating to 6,400 years ago, from Mummy Cave, an
archaeological site in northwestern Wyoming.
Key words: American marten, extinction, faunal associations, habitat preference, Martes americana, Martes americana
nobilis, Mummy Cave, noble marten, taxonomy
whether competition with the American marten was the cause
of its extinction (Grayson 1984, 1987; see also Heaton 1990).
The taxonomic status of the noble marten within the genus
Martes also has been debated. When Hall (1926) 1st described
the noble marten, he assigned it to subspecific status as Martes
caurina nobilis. Anderson (1970) assigned it to a separate
species, Martes nobilis, based on distinct morphometric
characteristics. More recently, the taxon was synonymized
with Martes americana by Youngman and Schueler (1991).
Subsequently, Anderson (1994) proposed it retain subspecific
status as M. americana nobilis.
The purpose of this paper is 2-fold. First, I introduce a new
Holocene find of a noble marten from Mummy Cave, an
archaeological site in northwestern Wyoming. This new
specimen, a right mandible, was recovered from a 6,400year-old stratum, and is the 1st Holocene occurrence of the
noble marten from Wyoming. Second, I analyze the fauna
associated with all noble marten finds and discuss the
implications for its adaptation, habitat preferences, extinction,
and taxonomic position.
Certain fauna, especially smaller taxa, are sensitive to local
environmental conditions and are useful paleoenvironmental
indicators (Barnosky et al. 2004; Lyman 1994; Mead and
Mead 1989). An analysis of climatically sensitive fauna consistently found in association with noble martens offers a
window into noble marten habitat preference that contributes
to our understanding of its extinction and taxonomic status.
For example, if boreal forest taxa dominate assemblages that
The cause of extinction of the noble marten (Martes
americana nobilis; also M. nobilis or M. caurina nobilis),
a late-Pleistocene true marten in western North America, and
its taxonomic position have been the subject of debate in recent
years. This large marten, 1st identified by Hall (1926), has been
identified from 17 localities in the western United States. Most
of these date to the late Pleistocene and, therefore, researchers
believed originally that this marten disappeared in the terminal
Pleistocene. More recently, specimens have been identified in
Holocene contexts as late as 3,000 years ago (Grayson 1984,
1987; Heaton 1990).
Because Martes americana (American marten—Turton
1806), a close relative of the noble marten, is a boreal forest
species, it was generally assumed that the noble marten also
was a boreal forest species (Anderson 1968; Kurtén and
Anderson 1980), and that competition with the American
marten may have caused its extinction (Anderson 1970; Kurtén
and Anderson 1980). American martens have been identified
from 3 sites along with noble martens; however, boreal fauna
or forest-adapted fauna are generally rare at Holocene find
sites, leading Grayson (1984, 1987) to suggest that the noble
marten had a more-generalized adaptation. He also questioned
* Correspondent: [email protected]
Ó 2009 American Society of Mammalogists
www.mammalogy.org
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February 2009
HUGHES—NOBLE MARTEN REVISITED
include noble martens and the geographic distribution of noble
martens is synonymous with that of American martens, it is
likely that the 2 martens were sympatric. This would support
the hypothesis that they competed with each other, perhaps
contributing to extinction of the noble marten. If habitat
preferences of the 2 taxa differ, this supports classifying them
into subspecies.
THE TAXONOMIC QUESTION
The taxonomic position of the noble marten within Martes
remains unclear. Mustelids 1st appeared in the Old World
during the early Miocene (Koepfli et al. 2008). Today, there are
3 subgenera of Martes (Martes, Pekania, and Charronia) and 8
living species, although recent genetic research suggests that
the fisher should be placed in its own genus, Pekania (Koepfli
et al. 2008). All of these taxa inhabit the boreal and temperate
regions of the Northern Hemisphere. The 5 species included in
the subgenus Martes are referred to as true martens and are
distinguished by the absence of an external median rootlet on
P4 and smaller size (Anderson 1970, 1994). Four species are
known as the Holarctic species group (M. zibellina [sable], M.
americana [American marten], M. martes [European pine
marten], and M. melampus [Japanese marten]) because they
share many similarities in morphology, behavior, and habitat
(Anderson 1994), and replace each other geographically across
the circumboreal region. Anderson (1970, 1994) suggested that
they be considered a ‘‘superspecies,’’ an allopatric group that is
too different to be included in a single species. The 5th species,
M. foina (stone marten), has long inhabited warm temperate
forests, meadows, and suburban areas in central and southern
Europe, and as a result has lost some of the adaptations of
martens to cold climates. In the northern parts of its range,
stone martens will interbreed with M. martes (Bakeyev 1994;
Proulx et al. 2004).
In addition to the noble marten, 3 species of Martes are
known from late-Pleistocene contexts in North America: M.
americana, a true marten, and 2 fishers (subgenus Pekania), M.
pennanti and M. diluviana (Kurtén and Anderson 1980; Powell
1981). Only M. americana and M. pennanti survive today.
Before 1953, 2 species of M. americana were recognized
in North America: M. americana (the eastern group) and M.
caurina (the western group—Anderson 1994; Hagmeier 1961).
Anderson (1970) noted that M. caurina has features more
similar to M. zibellina than to the eastern American marten, as
well as differences in shape of the skull and bullae, larger size,
and differences in size and shape of the upper molar.
Fossil evidence suggests that M. americana 1st entered
North America across Beringia sometime between 65,000 and
122,000 years ago. Anderson (1970) suggested that these
martens moved into eastern North America and were isolated
by the Laurentide ice sheet during the Pleistocene. She proposed that a later migration from Siberia populated the western
United States forming the caurina group. With the retreat of
glacial ice in the terminal Pleistocene, the American marten
expanded westward (Anderson 1970, 1994).
75
Wright (1953) and Hagmeier (1961) noted that the 2 species
intergrade in Montana and British Columbia, and they
proposed that the 2 be combined into 1 species. Since then,
caurina has been reduced to subspecific status (see Clark et al.
1987; Hall 1981; Wilson and Reeder 2005). Clark et al. (1987)
recognize 8 subspecies of M. americana that can be separated
into 2 subspecific groups, americana and caurina, based on
differences in cranial features and fossil history. Subspecies
previously identified as M. caurina are now subsumed under
M. americana caurina (Merriam 1890; vulpine—Rafinesque
1819; origenes—Rhoads 1902; sierra—Grinnell and Storer
1916; vancouverensis—Grinnell and Dixon 1926), martens of
the western United States. Two other subspecies, M. americana
humboldtensis (Grinnell and Dixon 1926) of Humboldt
County, California, and M. americana nesophila (Osgood
1901), inhabiting the islands west of British Columbia, Canada,
are included in the ‘‘caurina’’ group. The ‘‘americana’’ group is
represented by 5 subspecies distributed across the forested
regions of Canada, Alaska, and the eastern United States (Clark
et al. 1987).
Recent DNA evidence points to separate americana and
caurina clades (Carr and Hicks 1997; McGowan et al. 1999;
Stone and Cook 2002; Stone et al. 2002), stemming from
a single colonization event (Stone and Cook 2002). These 2
clades match the distribution of the 2 subspecific groups
defined by Clark et al. (1987). The caurina clade occurs in
western North America, extending from Admiralty Island in
southeastern Alaska south to Oregon and Wyoming (Carr and
Hicks 1997; Stone et al. 2002). The DNA evidence also points
to interbreeding between the 2 clades where they come into
contact, for example, 1 island in southeastern Alaska and
Montana (Stone and Cook 2002; Stone et al. 2002). According
to Stone et al. (2002), the DNA evidence is consistent with the
isolation of these 2 clades in glacial refugia during past glacial
periods and then a rapid late-Pleistocene spread of americana
westward as the ice retreated (Stone and Cook 2002).
The presence of a larger, late-Pleistocene marten was 1st
noted by Hall (1926) from specimens recovered from Samwell
and Potter Creek caves in northern California. These specimens
were intermediate in size between the modern American
marten and fisher and bore distinctive cranial and dental
features. He gave the specimens subspecific status as Martes
caurina nobilis (Hall 1926). Subsequently, he recognized that
these differences were not of taxonomic value and concluded
that the taxon did not differ sufficiently from the modern local
marten, Martes caurina sierrae (¼ M. americana sierrae), to
warrant subspecific recognition (Hall 1936).
Subsequent discoveries of the noble marten at Box Elder
Cave (Anderson 1968) and Jaguar Cave (Guilday and Adam
1967) prompted Anderson (1970) to revise the late-Pleistocene
and Holocene taxonomy of the mustelids. Using comprehensive morphometric comparisons of teeth and mandibles from
a large number of specimens, she assigned all 4 of the North
American Pleistocene martens and fishers to separate species,
M. americana, M. nobilis, M. diluviana, and M. pennanti.
According to Anderson (1970, 1994), what distinguished
the noble marten from other martens was its larger size
76
JOURNAL OF MAMMALOGY
(intermediate between a female fisher and male marten),
a broader skull, a more-robust P4 due to a narrower protocone,
and reduced indentation between the paracone and the
protocone. The trigonid on M1 is shorter than that of the other
martens, and on a plot of length of M1 versus length of the
trigonid, the noble marten lies on a different trend axis than do
the other 3 Pleistocene species of marten.
Youngman and Schueler (1991) conducted a discriminant
function analysis of caurina, americana, and nobilis tooth,
cranial, and humeri measurements, and concluded that size was
the most significant difference between the noble marten and
American marten, with the noble marten approximately 30%
larger. Metrically, the noble marten most closely approximated
the caurina group. Youngman and Schueler (1991) suggested
that the larger size of the noble marten resulted from
Pleistocene gigantism, and following Hall, proposed that it be
synonymized with M. americana (Youngman and Schueler
1991). Anderson (1994) pointed out that the noble marten
remains large throughout the Holocene and thus its larger size
is likely not the result of Pleistocene gigantism. She suggested
that the noble marten be given subspecific recognition as M.
americana nobilis to distinguish it from American martens
when found in prehistoric contexts (Anderson 1994). In light of
the taxonomic ambiguity and the reassignment of several previously identified americana specimens to noble martens since
1970, Graham and Graham (1994) recommend that the fossil
material be reevaluated. The subspecific designation proposed
by Anderson (1994), M. americana nobilis, is used to distinguish the noble marten from the American marten in this paper.
THE AMERICAN MARTEN
Because it has been assumed that noble martens maintained
adaptations similar to those of American martens, the diet,
habitat preference, and aspects of the physiology of American
martens are described here. The American marten is 1 of the 4
Holarctic martens, and like the others, has evolved physiological and behavioral adaptations to high-latitude, cold climates
(Ferguson and Larivière 2004). Some of these include low
population density, large home range for body size, long estrus,
delayed implantation, and a multimale mating system that
increases genetic variation where population densities are low
(Ferguson and Larivière 2004; Mead 1994; Ruggiero et al.
1994).
In general, population density scales to the 1.46 exponent
of body mass, but martens have such a low population density
that their density scales to one-tenth of this (Ruggiero et al.
1994). The home ranges of these solitary carnivores vary
between 1.1 and 8.1 km2. Several matings occur over a 30- to
45-day period in later summer, and often ovulation is not
stimulated until after the 1st mating (Clark and Stromberg
1987; Mead 1994; Ruggiero et al. 1994). Because of the large
size of home ranges and solitary nature of these mammals,
a female is unlikely to mate .3 times, and usually mates with
males from adjacent home ranges (Mead 1994). After mating,
female martens delay implantation to allow birthing between
mid-March and April of each year.
Vol. 90, No. 1
Today, American martens primarily inhabit the Hudsonian
and Canadian life zones in northern North America. These
zones are found at elevations above 2,000 m in the mountains
of western North America (Grinnell et al. 1937). The optimum
habitat for American martens is mature, old-growth, spruce–fir
communities with .30% canopy, a well-established understory
of fallen logs and stumps, and lush shrub and forb vegetation to
support microtine and sciurid prey (Clark et al. 1987; Martin
1994). American martens usually den in trees or logs (Ruggiero
et al. 1994: table 2). A dense forest understory is necessary in
winter to provide thermal protection, protection from predators,
and access to subnivean space where most winter prey are
captured (Martin 1994; Ruggiero et al. 1994).
Martens are opportunistic rodent generalists, with Myodes
(formerly Clethrionomys, forest species) and other voles as
a preferred food (Clark et al. 1987; Clevenger 1994; Martin
1994; Zalewski 2004). Diet breadth increases when abundances
of preferred prey are low, and then martens consume other
rodents, small rabbits, birds, insects, and plant material
(Clevenger 1994; Martin 1994; Zalewski 2004). Winter is the
limiting season and during this time of year, martens depend
more upon rodents than other types of food (Martin 1994;
Zalewski 2004).
American martens have important ecological relationships
with red squirrels (Tamiasciurus hudsonicus) and Douglas
squirrels (Tamiasciurus douglasii), both forest species, because
their middens are often used as resting sites (Ruggiero et al.
1994). In the forested regions of the subarctic, martens are most
abundant in areas with large, steep hills (Nelson 1973). In the
alpine regions of the Sierra Madres in California, American
martens have been observed using alpine talus and rock slides
adjacent to forests in summer (Grinnell et al. 1937). Riparian
areas may be used for resting or hunting (Ruggiero et al. 1994),
but a distance of more than 5 km of nonforested land below the
conifer zone is considered to be a complete barrier to marten
dispersal (Gibilisco 1994). In the montane West, local
extinctions occur because large basins separating forested
mountain islands prevent interbreeding between local populations (Gibilisco 1994).
Martens are vulnerable to habitat change and a reduction
in vole abundance (Buskirk and Powell 1994; Ferguson
and Larivière 2004; Martin 1994). As a result of human
expansion, hunting, and logging of forests, martens have
experienced a significant loss in range (Clark et al. 1987;
Gibilisco 1994).
THE MUMMY CAVE NOBLE MARTEN
Mummy Cave.— Mummy Cave is a multicomponent prehistoric human hunting site in the Absaroka Mountains of
northwestern Wyoming (Fig. 1; Hughes 2003; Husted and
Edgar 2002). Situated in a narrow mountain valley on the
eastern flank of the middle Rocky Mountains, the rock shelter
is 27 km east of Yellowstone National Park and 58 km west of
the semiarid Bighorn Basin. Erosion of the Absaroka volcanics
has created deep canyons and rugged, rocky topography. The
rock shelter was carved into a steep southwest-facing cliff by
February 2009
HUGHES—NOBLE MARTEN REVISITED
FIG. 1.—a) The location of Mummy Cave and other noble marten
(Martes americana nobilis) sites. Open symbols represent Holocene
sites; closed diamonds represent late-Pleistocene sites. Numbers
correspond to site names listed in Table 2 (data from Faunmap
Working Group 1994). Old Crow is not shown. b) The entrance to
Mummy Cave rock shelter is at the bottom center of photograph (Jack
Richards Photography, courtesy of the Buffalo Bill Historical Center,
Cody, Wyoming).
the North Fork of the Shoshone River during the late
Pleistocene (Moss 2002).
The present vegetation of northwestern Wyoming forms
distinct elevation zones (Baker 1986; Daubenmire 1943;
Gennett and Baker 1986; Waddington and Wright 1974;
Whitlock and Bartlein 1993). Shadscale (Atriplex confertifolia)
and sagebrush (Artemisia) dominate the semiarid Bighorn
Basin at 1,000–1,750 m above sea level. A sagebrush–
grassland belt (1,750–2,100 m) covers the foothills and lower
mountain valley terraces. From 2,000 to 2,800 m, a dense
coniferous forest flanks the mountains with Douglas-fir
(Pseudotsuga menziesii) and juniper (Juniperus) near the
bottom, lodgepole pine (Pinus contorta) throughout, and
subalpine fir (Abies lasiocarpa) and spruce (Picea) near upper
timberline. Above 2,800 m, timber gives way to alpine tundra.
Mummy Cave is presently located at the Douglas-fir and
foothills ecotone. Average annual temperature is between 38C
and 48C with average annual precipitation ranging from 76 to
77
102 cm/year (Hughes 2003). The rock shelter is adjacent to the
upper North Fork riparian community consisting of mountain
maple (Acer glabrum), cottonwood (Populus deltoides), cherry
(Prunus), service-berry (Amelanchier), willow (Salix), and
gooseberry (Ribes—Moss 2002).
Mummy Cave produced one of the most complete and welldated Holocene records of human occupation in the western
United States (Hughes 2003; Husted and Edgar 2002). More
than 38 different cultural strata spanning 9,500 years were
revealed in excavations and 22 of these produced faunal
materials (Hughes 2003). Bighorn sheep (Ovis canadensis)
dominate the faunal assemblages, but smaller mammals and
birds also are present. The mandible identified as M. americana
nobilis was recovered from stratum 12 dating to 6,400 6 75 SD
years ago (AA46120—Hughes 2003). An auditory bulla
indistinguishable from those of other American martens (M.
americana caurina) was recovered from stratum 6/7 with dates
ranging between 4,200 and 4,500 years ago (Hughes 2003).
The 4 taxa associated with the noble marten in stratum 12 are
abundant in the region today and reflect the rocky forest–foothills
ecotone where the site is presently located: O. canadensis, Lepus
americanus (snowshoe hare), Lepus townsendii (white-tailed
jackrabbit), and an unidentified grouse. Bighorn sheep prefer
open, rocky terrain and avoid forests (Wilson and Ruff 1999).
Snowshoe hares are most often found in coniferous forests
(Wilson and Ruff 1999), whereas white-tailed jackrabbits inhabit
grasslands (Lim 1987). The grouse could either be a ruffed
grouse, an inhabitant of conifer forests, or a sage grouse, an
inhabitant of sagebrush communities. Butchering marks on
these bones suggest that these taxa were acquired by human
hunters and transported to the rock shelter (Hughes 2003).
The mandible.— The Mummy Cave mandible (x5183) was
originally identified as M. americana by Arthur Harris in the
mid-1960s, well before Anderson (1994) published her revision
of the mustelids (Harris 2002). The mandible is now reassigned
to M. americana nobilis based on morphometric characteristics
of the dentition and mandible. The identification of this
specimen and that of the auditory bulla were confirmed
independently by Elaine Anderson in 2002 (E. Anderson, pers.
comm., April 2002).
The Mummy Cave specimen is a nearly complete right
mandible lacking only the incisors, canine, p1, and m2 (Figs. 2a
and 2b). The 2nd and 3rd premolars are broken, and the
protoconid and metaconid of the carnassial (m1) are damaged.
The Mummy Cave specimen fits well within the size range
of other noble martens (Table 1; data from Anderson 1970).
Three of the measurements, mandible length, height, and length
of the toothrow, reveal that the Mummy Cave mandible is
slightly smaller than the Pleistocene martens measured by
Anderson (1970), but the specimen is within the range of other
noble marten specimens.
The Mummy Cave carnassial is plotted with Anderson’s
(1970) marten data in Figs. 3 and 4. Length of the Mummy
Cave carnassial is closer to that of the modern and Pleistocene
fisher than to the American marten (Fig. 3). When length of the
m1 trigonid is plotted against the maximum length of the m1,
the Mummy Cave specimen fits well within the noble marten
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JOURNAL OF MAMMALOGY
Vol. 90, No. 1
FIG. 3.—Length measurements of m1 of 4 North American
Pleistocene martens (after Anderson 1970). The star marks the
position of the Mummy Cave specimen (x5183).
FIG. 2.—Noble marten (Martes americana nobilis) mandible from
Mummy Cave. a) Lingual view of mandible and toothrow, and b)
occlusal view of mandible revealing shape of carnassial. Scale is in
millimeters.
Fig. 1; Graham and Graham 1994; Kurtén and Anderson 1980).
Three of these sites have also produced American martens: Bell
Cave, Chimney Rock, and Snake Creek Cave. The known find
sites of the noble marten date to the late Pleistocene (n ¼ 13)
group (Fig. 4). This group is characterized by a different trend
axis suggesting that the trigonid is slightly shorter in noble
martens than in either pennanti or americana.
MATERIALS AND METHODS
With Mummy Cave added to the list, noble martens have
been identified from 18 sites in western North America (Table 2;
TABLE 1.—Measurements (in mm, 6 SD) of the noble marten
(Martes americana nobilis) mandible from Mummy Cave (central
column) compared against the range of all noble marten values
examined by Anderson (1970).
Measurement
x5183
Anderson 1970
Mandible length
Length of toothrow
m2 breadth
Mandible height
m1 length
m1 trigonid length
52.76
32.52
3.77
22.85
10.60
7.31
55.08 6 2.16 $
38.3 #, 34.96 6 1.33 $
25.27 6 0.82 $
11.08 6 0.19 #, 10.07 6 0.26 $
7.44 6 0.23 #, 6.93 6 0.20 $
FIG. 4.—Length of m1 plotted against length of m1 trigonid (after
Anderson 1970). The asterisk marks the position of the Mummy Cave
specimen (x5183).
February 2009
HUGHES—NOBLE MARTEN REVISITED
79
TABLE 2.—Noble marten (Martes americana nobilis) find localities with stratum and associated dates. The numbers in the left column refer to
site locations in Fig. 1a.
Map no.
Site
Stratum
Date (years ago)
1
Mummy Cave, Wyoming
12
6,400
2
Bell Cave, Wyoming
UN
12,240
3
Little Canyon Creek Cave, Wyoming
.10,170 6 250
4
Little Box Elder Cave, Wyoming
Below cultural
occupation
UN
.10,000
5
Natural Trap Cave, Wyoming
Glacial
14,67020,250
6
7
Chimney Rock, Colorado
North Cove, Nebraska
.24-inch depth
G
11,00013,000
12,96514,700
8
Jaguar Cave, Idaho
All strata behind seal
10,00012,000
9
12
3,270 6 110
10
11
12
Dry Creek Rockshelter, Idaho
(stratum 12)
Moonshiner Carnivore Trap, Idaho
Wilson Butte Cave, Idaho
Hidden Cave, Nevada
UN
C (lower)
II; IXXIII
13
14
15
Smith Creek Cave, Nevada
Snake Creek Burial Cave, Nevada
Bronco Charlie Cave, Nevada
Reddish brown silt
III
Unit 2S, 245250 cm
.5,175
Approximately 10,000
II ¼ 3,6003,700;
IXXIII ¼ .7,500
.12,000
9,46015,100
3,500
16
Potter Cave, California
UN
Late Pleistocene—older
17
Samwel Cave, California
Chamber 2
Late Pleistocene—younger
18
Old Crow, Yukon, Canada
Localities 11A, 28, 65
Late Pleistocene
and Holocene (n ¼ 5) with the most recent dates at 3,000 years
ago (Table 2). Most of the late-Pleistocene sites are natural
deposits, whereas most of the Holocene sites are cultural
deposits (Table 3). Of the 3 types of sites represented, animal
traps, natural deposits, and cultural deposits, the first 2 likely
offer less-biased samples of the fauna living in the area,
whereas humans may have transported smaller prey some
distance to their campsite. Unfortunately, no information exists
on which fauna, if any, were brought in by predators or what
distance they might have been transported at most of these
sites, but if faunal patterns in natural and cultural sites differ
markedly from those in natural traps, it would suggest that
predation practices may have biased samples.
From Table 2 it appears that most of the Pleistocene
noble marten find sites, especially in the interior West, date to
the terminal Pleistocene, 10,000 and 14,000 years ago. The
specimens from Potter Creek and Samwel Caves in California
are associated with full glacial fauna (Furlong 1906), and may
be the earliest occurrences of the noble marten.
Sites are found in the foothills and mountain valleys of the
Rocky Mountains in Wyoming and Colorado, the mountain
edges and foothills ringing the northern and western edges of
the Great Basin, and in the foothills of northern California with
2 outliers, 1 in Nebraska’s shortgrass prairie (North Cove), and
Reference
Harris 2002; Hughes 2003;
Husted and Edgar 2002
Walker 1987; Zeimens and
Walker 1974
Shaw 1980; Walker 1987
Anderson 1968; Hager 1972;
Walker 1987
Chomko and Gilbert 1987;
Walker 1987
Anderson 1974; Hager 1972
Faunmap Working Group 1994;
Stewart 1987
Guilday and Adam 1967;
Kurtén and Anderson 1972
Webster 1978
White et al. 1984
Gruhn 1961; Kurtén and Anderson 1980
Grayson 1984, 1985; Thomas 1985
Mead et al. 1982; Miller 1979
Heaton 1990; Mead and Mead 1989
Casjens 1974; Grayson 1984, 1987;
Mead and Mead 1989; Speiss 1974
Faunmap Working Group 1994;
Hall 1926; Kurtèn and Anderson
1980; Sinclair 1903
Furlong 1906; Hall 1926; Kurtèn and
Anderson 1980
Harington 1977; Kurtèn and
Anderson 1980
the 2nd in Old Crow Basin, Yukon, Canada (Table 3; Fig. 1).
The northern Californian foothills have a milder climate and
support a more-mesic type of vegetation and fauna than the
interior West. Old Crow and North Cove occur at significantly
lower elevations than the other sites (Table 3). The Old Crow
specimens were recovered from redeposited channel sediments
along the Old Crow River and depositional context and faunal
associations are not known. The North Cove specimen was
recovered with other late-Pleistocene fauna on the north edge
of the Republican River floodplain. Most sites are located in
open rocky canyons adjacent to river or stream channels (Table
3). Those sites where no permanent water is present today
include Snake Creek Burial Cave, Jaguar Cave, Moonshiner
Carnivore Trap, and Wilson Butte Cave.
Fauna recovered from the find sites are compared in 2 ways:
1st by life-zone association, and 2nd by habitat preference
(Appendix I). The life-zone concept, originally proposed by
Merriam (1894), is too simplistic to accurately describe
variation in local plant and animal communities (Ingles 1965;
Odum 1971), but it has the advantage of allowing a comparison
of fauna and vegetation across a regional scale (Ingles 1965).
For this paper, Merriam’s (1894) basic life-zone concept is
modified to distinguish cool-adapted, mesic fauna from morexeric fauna. The original 6 zones are compressed into 3
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JOURNAL OF MAMMALOGY
TABLE 3.—Present environmental characteristics of noble marten (Martes americana nobilis) find sites (references are listed in Table 2).
Site
Late Pleistocene
Little Canyon Creek Cave,
Wyoming
Elevation (m)
Deposit
1,399
Natural
Bell Cave, Wyoming
2,379
Natural
Little Box Elder Cave,
Wyoming
1,677
Natural
Natural Trap Cave, Wyoming
1,510
Natural trap
Chimney Rock Animal Trap,
Colorado
2,410
Natural trap
Smith Creek Cave, Nevada
1,878
Natural
455
Natural
Samwel Cave, California
Snake Creek Burial Cave,
Nevada
1,731
Natural trap
Jaguar Cave, Idaho
2,257
Cultural
595
Natural
1,344
Cultural
380
Natural
Mummy Cave, Wyoming
1,950
Cultural
Moonshiner Carnivore Trap, Idaho
1,480
North Cove, Nebraska
Wilson Butte Cave, Idaho
Old Crow, Yukon, Canada
Location
River canyon in Bighorn
Mountains foothills;
permanent stream
River canyon in Laramie
Mountains foothills;
ephemeral stream
River canyon in Laramie
Mountains foothills;
permanent stream
Karst sinkhole on Little
Mountain, Bighorn
Mountains; ephemeral
stream nearby
Mountain foothills;
permanent stream
at 0.4 km
River canyon in foothills;
ephemeral stream
River canyon in Cascade
Range foothills;
permanent river
Sinkhole on bajada
developed from Snake
Creek; no water
source near
Riverine canyon in
foothills of Beaverhead
Mountains; dry
streambed
Tributary of Republican
River; permanent river
Lava blister on Snake
River Plain; no
known water source
Displaced; location
unknown
Modern environment
Modern vegetation
Open, rocky, semiarid
Transitional (sagebrushjuniper)
with riparian community
Open, rocky, semiarid
Mountain mahogany, big sage,
grass with pine forest at 2 km
Open, rocky, semiarid
Mountain mahogany, big sage,
grass community with isolated
stands of timber pine at 2 km,
riparian at 1.5 km, and
shortgrass prairie at 1 km
Transitional (sagebrushmountain
mahoganylimber pine)
Open, cliffs, semiarid
Open, cliffs
Transitional with Canadian zone
12.8 km SW
Talus slopes, semiarid
Cliffs
Pinyonjuniper forest with
shadscale and sagebrush
communities below and conifer
forest at higher elevations and
north slopes
Transitional and riparian
Open, rocky, arid
Greasewoodshadscale
Open, rocky, semiarid
Upper Sonoran (grasssage
with coniferous forest at
higher elevations)
Unknown
Prairiesteppe on benches;
wooded riparian along
river floodplain
Upper Sonoran
Open, rocky, semiarid
Tundra
Tundra with spruce, willow,
and alder along banks and
meander scars
River canyon in
Absaroka Mountains;
permanent river
Partially open, cliffs,
semiarid
Natural
Lava blister on Snake
River Plain; no
known water source
Open, rocky, semiarid
Transitional
(sagebrushgrassland
Douglas-fir forest ecotone)
with riparian
Upper Sonoran
960
Cultural
Riverine canyon
in foothills;
permanent stream
Rocky, open, semiarid
Hidden Cave, Idaho
1,251
Cultural
Rocky, open, arid
Bronco Charlie Cave,
Nevada
2,134
Cultural
River canyon in foothills
of Stillwater Mountains;
dry streambed
River canyon in Ruby
Mountains;
dry streambed
Early Holocene
Later Holocene
Dry Creek Cave, Idaho
Cliffs, talus, open
Bluebunch grasssagebrush
with riparian in canyon and
ponderosa pine and Douglas-fir
forest at higher elevations
Lower Sonoran with
pinyonjuniper at higher
elevations
Upper Sonoran (pinyonjuniper
and sagebrush)
February 2009
HUGHES—NOBLE MARTEN REVISITED
81
FIG. 5.—Distribution of late-Pleistocene and Holocene noble
marten (Martes americana nobilis) and American marten (Martes
americana) fossil localities overlain by the modern distribution
(hatched area) and the historic distribution (dark line) of the American
marten. The dashed line reflects the dividing point between the
caurina and americana clades (map adapted from Graham and
Graham [1994: figure 2.2]; see also Gibilisco [1994: figure 3.4]).
categories: Alpine (Tundra), Boreal (Hudsonian–Canadian),
and Sonoran (Transitional, Upper, and Lower Sonoran).
Tundra species are those adapted to arctic tundra environments, which are cool and dry. Only 1 species found with noble
martens is assigned to this group, Dicrostonyx (collared
lemming). The closest analogs to arctic tundra are highelevation alpine meadows, but most of the fauna that inhabit
these also use subalpine meadows in the Hudsonian–Canadian
zones, and thus, are assigned to the latter group.
The fauna included in the Hudsonian–Canadian zone are
referred to as montane, boreal, or mesic species in the literature
(Graham et al. 1987; Grayson 1977, 1987, 1993; Heaton 1990;
Lundelius et al. 1983; Walker 1987), and have northerly
distributions. Throughout the rest of this paper, the term
‘‘boreal’’ is used to identify this group.
The term ‘‘boreal’’ has several meanings in the literature. In
its true meaning, it refers to the expansive spruce, fir, hemlock,
and larch forests of Canada (the taiga), but the term is often
expanded to include the mesic, montane conifer forests of
western North America. Others use boreal to refer to cooladapted species (Lundelius et al. 1983) or mesic species
(Grayson 1977, 1993), for example, those species requiring
more-moist environments. Boreal is used here to identify taxa
FIG. 6.—Percentage of Pleistocene and Holocene mammals found
with noble martens (Martes americana nobilis) at all sites associated
with 1 of the 3 life zones (Sonoran, Boreal, and Tundra) defined in
text. Percentages are rounded to nearest whole percent.
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Vol. 90, No. 1
February 2009
HUGHES—NOBLE MARTEN REVISITED
that are adapted to cold, moist climates, but not necessarily
forested environments. Cool, mesic environments were more
widespread during the late Pleistocene (Graham and Lundelius
1984; Grayson 1993; Thompson et al. 1993), but today are
associated with northerly regions, dense forests, and highelevation environments.
The taxa assigned to the Sonoran group are those generally
associated with the transitional, Upper and Lower Sonoran life
zones today, fauna better adapted to warm, xeric conditions. In
the intermountain West, these taxa inhabit the desert basins up
to the sagebrush–pinyon–juniper zones that flank the base of
the mountains (Baker 1983; Clark and Stromberg 1987; Heaton
1990). In northern California, the transitional life zone is
forested with yellow pine, sugar pine, Douglas-fir, white fir,
cedar, spruce, and redwood. The Upper Sonoran is characterized by open deciduous forest with saltbush, sage, and
rabbitbrush (Ingles 1965), and the Lower Sonoran with yucca,
valley oak, agave, and many kinds of cacti (Ingles 1965). Some
taxa associated with the noble marten, such as Mephitis
mephitis (striped skunk) or Mustela frenata (long-tailed weasel),
do not fit easily into any 1 of these categories, and were not
assigned to a life zone.
Each life zone is characterized by a variety of habitats or
microenvironments. Within the montane zone, for example,
there are closed-canopied forests, alpine and subalpine
meadows, wet or dry meadows, parklands, riparian communities, and dry, south-facing rocky escarpments with minimal
vegetation. Habitat associations were assigned to each taxon
through a review of the following sources: Wilson and Ruff
(1999), Clark and Stromberg (1987), Ingles (1965), and various
issues of Mammalian Species (Carroll and Genoways 1980;
Chapman 1975; Clark et al. 1987; Eshelman and Sonnemann
2000; Hillman and Clark 1980; Jones and Baxter 2004; King
1983; Larivière and Pasitschniak-Arts 1996; Lim 1987;
Michener and Koeppl 1985; Poglayen-Neuwall and Toweill
1988; Powell 1981; Rickart 1987; Sheffield and Thomas 1997;
Verts and Carraway 1999; Wade-Smith and Verts 1982; Zegers
1984).
Habitat associations for each taxon are listed in Appendix I.
Species that live in or use trees or are adapted to closedcanopied forested environments are indicated by ‘‘F.’’ Open
(O) refers to any type of open vegetative habitat including
mountain meadows, alpine tundra, and grasslands, sagebrush,
although the 1 tundra species is indicated by ‘‘T.’’ Edge (E)
refers to woodland–grassland edges. Riparian (R) includes all
types of wetlands including rivers, streams, lakes, ponds, and
marshes. Many riparian communities consist of small stands of
trees and a dense understory of shrubs. Species assigned to
rocky habitats (X) either nest in caves, such as Neotoma
cinerea (bushy-tailed woodrat), or use rocks and talus slopes
for nesting, hunting, or both. Meadow species (M) are those
83
that are generally adapted to more-mesic herbaceous meadows
characteristic of alpine and montane settings. Sagebrush (S) is a
common vegetation type of the Upper Sonoran and transitional
life zones today. Grassland (G) taxa are those adapted to dense
shortgrass prairie characteristic of the western Great Plains.
Some taxa also prefer habitats with a dense understory of
shrubs for hunting and nesting, and these preferences are
indicated by (U).
Taxa used in this analysis include climatically sensitive
fauna found in the same stratum as noble martens in 15 of the
find sites (Appendix I). Only taxa appearing in .1 site and
only sites with a sample of 3 taxa are included in the analysis.
Three sites are excluded because either the deposit was
secondary (Old Crow) or ,3 small to medium-sized taxa were
associated with the noble marten (North Cove and Dry Creek
Cave). Some of the strata represent longer periods of deposition
than others and these are generally represented by a larger
sample size.
Hidden Cave, Nevada, produced noble marten remains from
both late-Pleistocene and late-Holocene strata. The Holocene
occurrence is a single M1, and although Grayson cautions that
this specimen might be displaced from earlier strata (Grayson
1985), this stratum is included because no direct evidence of
displacement is given (Grayson 1985). The 2 Hidden Cave
occurrences are treated separately. In Appendix I, the
Pleistocene stratum at Hidden Cave is labeled ‘‘P,’’ the
Holocene stratum, ‘‘H.’’
RESULTS
Life-zone association.— Approximately 75% of the taxa
reported from noble marten find sites in both Plesitocene and
Holocene strata are boreal species, or those adapted to cooler,
mesic conditions (Fig. 6). The only significant difference
between the Pleistocene and Holocene fauna is the disappearance of Dicrostonyx in the Holocene. The dominance of mesic
and cool-adapted fauna in association with noble martens
suggests that this marten was likewise adapted to cool and
mesic environments.
Habitat preference.— Habitat preferences are condensed into
5 groups: woodland–forest, open (grasslands, sagebrush, and
meadow), edge, riparian, and rocky. The mix and percentage of
fauna is fairly consistent across most of these sites (Fig. 7).
Open-adapted species clearly dominate all but 2 of the
assemblages and average 52.4% (SD ¼ 14%) of fauna. The 2
exceptions are the assemblages from Samwel and Mummy
caves, where open and forest species are represented equally.
Only 3 taxa are associated with the noble marten at Mummy
Cave, results that could be influenced by the small sample size.
Seven taxa are represented at Samwel Cave and a moreforested environment is suggested.
FIG. 7.—Percentage of mammals found with noble martens (Martes americana nobilis) associated with 1 of 5 defined habitat types: woodland
(forest), open, edge, riparian, or rocky, at each noble marten find site. Percentages are rounded to nearest whole percent. Pleistocene and Holocene
faunal assemblages are identified by (P) and (H), respectively.
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JOURNAL OF MAMMALOGY
Rock-loving species are present at all sites and form the next
largest group of associated fauna (22.2%, SD ¼ 13.5%). Forest
species comprise 11.2% (SD ¼ 10.4%) of fauna, suggesting
a nearby woodland community. Small numbers of edge (7.2%,
SD ¼ 6.7%) and riparian species (7.0%, SD ¼ 7.2%) further
support this. Riparian species are present in 10 of the faunal
assemblages and may point to a preference for riparian
communities. When combined, the faunal associations suggest
that the noble marten preferred open, rocky habitats near
a woodland–riparian community.
Distribution.— If the noble marten was sympatric with the
American marten, its distribution should mirror that of the
American marten. The western range of the American marten is
synchronous with the mountain regions, which are today
flanked by dense coniferous forest, the primary habitat of the
American marten. According to Graham and Graham (1994),
late-Pleistocene and early-Holocene occurrences of the noble
marten are within the historic range of M. americana, whereas
3 late-Holocene sites, Dry Creek, Bronco Charlie, and Hidden
Cave, are outside of its historic range. Noble marten sites are
located along the periphery of American marten historic and
modern range, but rarely in the interior (Fig. 5). This might
suggest that noble martens preferred lower-elevation foothills
whereas the American marten inhabited higher-elevation
montane forests. Descriptions of the modern landscape and
vegetation communities associated with noble marten sites
support this; most of the noble marten sites are located in river
valleys in the montane foothills (Table 3).
If the noble marten inhabited foothill environments and the
American marten inhabited forested upper mountain zones,
then elevation records of find locations should demonstrate that
the American marten existed at higher elevations where their
distributions overlap. Pleistocene and Holocene records of findsite elevations for the 2 subspecies are compared to determine
if an elevational difference does exist (Fig. 8). The modern
American marten sample consists of martens that were trapped
at known elevations and are listed in Grinnell et al. (1937), Hall
(1946), and Long (1965). Although there are numerous
elevation records for recent American martens, there are very
few late-Pleistocene records, as noted by Graham and Graham
(1994), making a temporal comparison difficult. Only 4 fossil
localities could be identified with confirmed Pleistocene
records of M. americana. Three of these also yielded specimens of the noble marten, although at 2 of these sites, Chimney
Rock and Bell Cave, noble martens appear to be stratigraphically below modern martens (Anderson 1974; Hager
1972). Although they are included in the Pleistocene sample,
these American martens also may date to the Holocene. The 4th
Pleistocene record is Crystal Ball Cave (Heaton 1988).
The elevation differences between late-Pleistocene records
of noble and American martens are not statistically significant,
as might be expected when 3 out of 4 of these sites contained
both subspecies (t ¼ 1.756, P ¼ 0.098; n ¼ 4 americana, n ¼
14 nobilis; Fig. 8). However, the Holocene group, represented
by a much larger sample of American martens, demonstrates
a significant difference in elevation (t ¼ 4.505, P , 0.0001,
n ¼ 31 americana, n ¼ 5 nobilis; Fig. 8), with the American
FIG. 8.—Mean elevation records of Pleistocene and Holocene
martens. Bars represent a 95% confidence interval.
marten generally found above 2,000 m and the noble marten
below 2,000 m. Two of the 3 Pleistocene sites that contained
both noble and American martens have elevations . 2,000 m
(Bell Cave and Chimney Rock). Examination of these data
suggests that at least in the Holocene, noble martens generally
inhabited elevations below those of American martens.
DISCUSSION
Adaptation and habitat preference.— Although the sample
of noble marten sites is small, a consistent pattern emerges
from the faunal associations. Both Pleistocene and Holocene
noble martens are associated with fauna adapted to mesic,
open environments with small numbers of riparian, forest, and
February 2009
HUGHES—NOBLE MARTEN REVISITED
rock-loving taxa present. Forest species make up only a small
percentage of the fauna at these sites. This would suggest that
the noble marten is not a boreal species when ‘‘boreal’’ refers to
a northern forest habitat. The noble marten can be considered
a boreal species when boreal refers to a cool, mesic climate.
The meaning attributed to boreal has contributed to a misunderstanding of noble marten habitat and extinction.
Species that routinely inhabit open landscapes dominate the
noble marten faunal assemblages, and reflect a mix of moist
meadows (e.g., Microtus montanus [montane vole]), shortgrass
or tallgrass prairies (e.g., Spermophilus richardsonii [Richardson’s ground squirrel] and Mustela nigripes [black-footed
ferret]), dense sagebrush (e.g., Lemmiscus curtatus [sagebrush
vole]), and dry sagebrush (e.g., Spermophilus mollis [Piute
ground squirrel]). The fauna associated with the noble marten
at the 2 sites not included in this analysis, North Cove,
Nebraska, and Dry Creek, Idaho, also reflect a more-open
environment. Blarina brevicauda (northern short-tailed shrew)
and Bison bison (American bison) from North Cove are found
in prairie environments (Faunmap Working Group 1994;
Lundelius et al. 1983). O. canadensis (bighorn sheep) and
Antilocapra (pronghorn), both recovered from Dry Creek
Rockshelter, inhabit sagebrush–grasslands. The variety of open
habitats represented in the noble marten assemblages may
reflect the mix of fauna coexisting in late-Pleistocene open
environments.
Taxa that inhabit rocky landscapes are present in small
numbers in the noble marten assemblages. This might be
expected given that most noble marten remains have been
recovered from caves or rock shelters, but also suggests that
noble martens were common in these habitats.
Small numbers of woodland, riparian, and edge species form
a background to the open-adapted species in the assemblages.
Because riparian species turn up frequently and many of the
noble marten find sites are located in river valleys, an
association with wet habitats is suggested. Riparian species
also are present in 2 Holocene assemblages where permanent
water sources do not exist today, Moonshiner Carnivore Trap
and Hidden Cave stratum 3. Moonshiner Carnivore Trap is
a lava blister on the Snake River Plain far from water today, but
White et al. (1984) note that snow building up in the bottom of
the trap creates a cool, moist environment. During the early
Holocene, water tables in the Snake River Plain were higher
(Davis et al. 1986), and the potential existed for seeps or ponds
in Moonshiner Cave. Stratum 3 at Hidden Cave revealed pollen
and seeds from riparian plants probably derived from a marsh
that existed 3,000 years ago below the cave entrance (Wigand
and Mehringer 1985).
The presence of both woodland and riparian species might
suggest that the noble marten inhabited open areas at the edges
of wooded riverine communities. These communities are
characterized by a dense understory of shrubs that offer
protection and an abundance of microtine and other small prey.
Some of the rodents and lagomorphs in the faunal assemblages,
for example, Spermophilus lateralis (golden-mantled ground
squirrel, n ¼ 7), Phenacomys intermedius (western heather
vole, n ¼ 4), L. americanus (snowshoe hare, n ¼ 4), and
85
Sylvilagus audubonii (desert cottontail, n ¼ 2), prefer a dense
understory. Edge communities also attract other carnivores
such as Vulpes vulpes (red fox, n ¼ 9), Mustela erminea
(ermine, n ¼ 5), Mustela frenata (long-tailed weasel, n ¼ 9),
and Neovison vison (American mink, n ¼ 5; see Appendix I),
all of which frequently appear in noble marten assemblages.
The pattern of faunal associations remains consistent
throughout the terminal Pleistocene and Holocene, even though
environments changed significantly during these periods. The
cool and mesic terminal Pleistocene environment became
increasingly more xeric through the early and middle Holocene
(Beiswenger 1991; Davis et al. 1986; Grayson 1993;
Thompson et al. 1993; Whitlock 1993), yet noble martens
continued to select mesic habitats locally. In the Holocene,
lower-elevation mesic sites are often associated with riparian
communities (Buskirk and Powell 1994), including riverine
valleys, springs, swamps, ponds, and other wetlands. The
association of the noble marten with riparian communities may
have become increasingly important in the Holocene as the
climate became more arid, although terminal Pleistocene sites
are frequently near water sources as well.
The difference in habitat between the noble and American
marten suggests that the 2 were not sympatric. Both were
adapted to cool, mesic climates, but whereas the American
marten was adapted to the northern taiga, or northern forests
commonly associated with boreal conditions, the noble marten
seems to have preferred open, rocky foothills near the edges of
forested riparian communities. As an inhabitant of lowerelevation sites, the noble marten was likely more tolerant of
warmer temperatures than the American marten, and may have
evolved in a warm, mesic environment, perhaps at lower
elevations, in the late Pleistocene. With so few late-Pleistocene
American marten specimens in the West, it is not clear what its
preferences were during the late Pleistocene, but with an
adaptation to mesic, closed-canopied forest, this taxon may
have inhabited lower-elevation forests during the Pleistocene
and followed these to higher elevations in the Holocene.
The extinction question.— If the noble marten and the
American marten were not sympatric, then competition
between the 2 subspecies was not the cause of extinction of
the noble marten as suggested by Anderson (1970) and Kurtén
and Anderson (1980). After surviving the Pleistocene–
Holocene transition, many researchers have wondered what
caused its extinction well into the Holocene (Graham and
Graham 1994; Grayson 1984, 1987; Heaton 1990). Graham
and Graham (1994) propose that habitat fragmentation caused
by Holocene climate change, a factor in the disappearance of
other small to medium-sized taxa in parts of the Great Basin
(Grayson 1987), may be the primary cause of its extinction.
Barnosky et al. (2004) noted that small mammals are more
affected by climatic warming than are larger animals. With the
small number of Pleistocene and Holocene records of the 2
martens in the American West, it is unlikely that this question
will be answered soon, but several facts relevant to its
extinction are presented below.
Certain cranial and dental differences distinguish the noble
marten from the American marten (Anderson 1970; Youngman
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and Schueler 1991). Based on its morphological characteristics
(Youngman and Schueler 1991) and its distribution in western
North American, it is more similar to the subspecific caurina
clade of M. americana. At some point in the late Pleistocene,
the noble marten was reproductively isolated from other
members of the clade and evolved minor morphological and
behavioral differences including larger body size and an
adaptation to mesic, open, rocky foothill environments,
environments that were likely more widespread in the late
Pleistocene.
Hall (1926) noted that the noble martens in California bear
a closer resemblance to M. americana caurina (¼ sierrae),
martens of the upper-elevation Cascade Range, than to M.
americana humboldtensis (¼ M. caurina humboldtensis), a rare
subspecies, allopatric with caurina, inhabiting the lowerelevation northern coastal ranges of California (Grinnell et al.
1937). The martens of the Cascade Range have since been
conjoined with the martens of the Pacific Coast ranges to the
north. According to Hall (1926), the noble marten most closely
resembles atypical specimens of M. americana caurina (¼
sierrae) recovered from Trinity and Siskiyou counties in
northern California. Hall (1926) notes that these atypical
specimens are regarded as intergrades between the martens of
the Sierra Nevada Mountains and the Pacific Coast range to the
north, and inhabit the lower-elevation Trinity, Scott, and
Salmon mountains in north-central California west of Shasta
County where Samwel and Potter Creek caves are located. Hall
(1926:130) wrote: ‘‘Judging from the morphologic characters
presented by the Pleistocene form [noble marten], it is clearly
related to the martens inhabiting north-central California to-day
and might well be regarded as belonging in or near the group
from which the modern forms of the region have taken their
origin. The presence of living martens in the region of the cave
deposit [Samwel and Potter Creek caves] coupled with the fact
that only slight structural differences exist between the Recent
and Pleistocene forms strongly suggest such relationship.’’
Most noble martens found at interior sites date to the
terminal Pleistocene, suggesting a possible expansion of the
range of the noble marten at this time. The noble marten may
have its origin in the mesic foothills of northern California
during the last glacial advance. Glaciers in the high mountains
of California may have isolated groups of martens in lowerelevation, forested river valleys where increased competition
may have allowed some to evolve a suite of behaviorial and
physiologic adaptations enabling them to hunt in open, mesic
foothills and riparian communities during the late Pleistocene.
Environmental evidence suggests that the western flanks of the
Sierras were more open during the late Pleistocene (Thompson
et al. 1993), and this evidence is supported by high percentages
of open-adapted fauna recovered from Samwel and Potter
Creek caves.
Terminal Pleistocene environmental conditions in the interior West were likely favorable to noble marten dispersal
because the climate was cool and moist, and mesic meadows
and parklands existed in the western basins (Grayson 1993;
Madsen and Currey 1979; Mead et al. 1982). Corridors
eastward may have opened with the retreat of glacial ice around
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14,000 years ago. In the Great Basin, increased moisture over
the next 2,000 years filled the valley bottoms with lakes (Lake
Bonneville, Lake Lahontan, and others), surrounded by
sagebrush steppe (Grayson 1993; Mead et al. 1982). In
northwest Wyoming, herb and shrub communities invaded
the deglaciated landscape around 14,000 years ago (Whitlock
1993; Whitlock and Bartlein 1993), and in southern Idaho,
Artemisia (sagebrush) steppe dominated the Snake River Plain
throughout the glacial and early postglacial period (Beiswenger
1991).
With increasing aridity after 10,000 years ago, mesic habitats
slowly disappeared in the montane basins and foothills. Great
Basin lakes contracted to marshes and shallow ponds (Grayson
1993; Thompson et al. 1993). The mesic forests of Idaho and
Wyoming migrated to higher elevations on mountainsides
and mesic meadows and Artemisia (sagebrush) steppe were
replaced by a semidesert steppe of Artemisia and cheno-ams
(chenopodium–amaranth species—Beiswenger 1991). Around
9,500 years ago, the hardy and fire-adapted Pinus contorta
(lodgepole pine) invaded all mountainous bedrock substrates
below the subalpine forest forming the core of the closedcanopied coniferous forest of the Rocky Mountains that exists
today (Knight 1994; Madsen and Currey 1979; Whitlock 1993;
Thompson et al. 1993). The noble marten, a species that was
adapted to open environments and cool, mesic conditions,
could become isolated and locally extinct as mesic environments disappeared in the foothills and basins.
That so few late-Pleistocene occurrences of the American
marten exist in the interior American West is odd. The
environment may not have been favorable for supporting large
numbers at that time, or they may have become isolated in the
eastern forests and western coastal mountains during the last
glacial advance. It may also be a sampling problem in that this
subspecies rarely used rock shelters or caves where the best
samples of fossil material are preserved.
Human hunting may have contributed to the extinction of the
noble marten, as suggested by Graham and Graham (1994).
Both noble and American martens are found in archaeological
assemblages from later-Holocene sites (see Graham and
Graham [1994] for a list of these sites). The American marten
was one of the most valuable fur-bearing animals in the recent
past and continues to be so today (Buskirk 1994; Clark et al.
1987; Grinnell et al. 1937; Mandelbaum 1979; Nelson 1973).
During the American fur trade, the marten was 2nd only to
beavers in pelts trapped and dollars earned (Clark et al. 1987;
Clark and Stromberg 1987).
Some of the same adaptations to northern boreal forest
environments make martens vulnerable to extinction in morefragmented environments: low population densities, large
home ranges for body size, small litters, and multimale mating
systems (Buskirk 1994; Ferguson and Larivière 2004;
Ruggiero et al. 1994). Multimale mating systems promote
sexual selection and increase genetic variation where population densities are low, but this reproductive strategy requires
gene flow between populations or local extinctions will occur
(Ferguson and Larivière 2004). Low population densities make
it more difficult for martens to find mates, especially in
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HUGHES—NOBLE MARTEN REVISITED
a multimale system of reproduction. Thus, a loss of habitat or
habitat isolation could significantly affect the ability of martens
to reproduce (Buskirk and Powell 1994; Martin 1994). With
the onset of more-xeric conditions in the Holocene, lowerelevation mesic environments decreased and marten populations may have become isolated in more-favorable locations.
Species with low population densities also are sensitive to
fluctuations in prey numbers (Clark et al. 1987; Martin 1994;
Powell 1994; Zalewski 2004), and this could lead to local
extinctions. If increasing aridity in the later Holocene reduced
prey richness and diversity, competition with other carnivores
such as M. frenata, N. vison, or V. vulpes, all adapted to similar
environments as the noble marten, may have contributed to
noble marten extinction.
As discussed above, a number of factors could have
contributed to the gradual extinction of the noble marten in
the late Holocene, including increasing aridity, loss of lowelevation mesic habitats, habitat isolation, reduction of prey,
overhunting, increased competition with other carnivores,
marten reproductive practices, and low population densities.
The precise cause of noble marten extinction remains uncertain,
but the particular physiological and behavioral traits of this
taxon and its apparent adaptation to open, mesic foothill
environments make it vulnerable to extinction in an increasingly arid Holocene environment, as proposed by Graham
and Graham (1994).
If americana and nobilis were not sympatric, as this research
suggests, the noble marten may indeed be a separate subspecies
as Anderson (1994) proposes. However, given the morphological similarity between the 2 subspecies, the reassignment of
several previously identified americana specimens to nobilis,
and the poor representation of the American marten in latePleistocene contexts in the West, the remains of the 2
subspecies should be reevaluated to settle this question once
and for all.
ACKNOWLEDGMENTS
I am deeply grateful to the Buffalo Bill Historical Center in Cody,
Wyoming, which not only gave me access to the Mummy Cave faunal
assemblage for study, but also provided other kinds of support. I also
am grateful to Elaine Anderson, who confirmed the identification of
the Mummy Cave specimens shortly before her death in April 2002.
The Mummy Cave mandible will always be linked to her memory, and
this paper is dedicated to Elaine’s exceptional contribution to
mammalogy and mustelid taxonomy. I am grateful to D. Grayson
for his initial identification of the specimen and to U. Albarella, D.
Walker, T. Heaton, K. Bovy, and several anonymous reviewers who
read drafts of this paper. This paper is part of a larger project funded
by the Buffalo Bill Historical Center and National Science Foundation
Dissertation Improvement Grant 9905628. No research was conducted
on live animals.
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February 2009
HUGHES—NOBLE MARTEN REVISITED
91
APPENDIX I
Small taxa associated with noble martens (Martes americana nobilis; compiled from references in Table 2; see ‘‘Materials and Methods’’
for habitat definitions). The number of specimens of noble martens is given when multiple specimens were reported; other taxa and single
records of noble martens are indicated by presence (þ) or absence (blank). Total number of taxa reported for each site is given at the bottom of
each column.
Wyomingb
Taxa
Coloradoc
Habitata NT LBE BC LCC MC
M. a. nobilis
CR
Idahod
Nevadae
WB JC MSCT SCC SNC HCP HCH BRC
þ
5
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
47
Californiaf
þ
46
25
1
þ
1
1
1
PC
SWL
þ
þ
Total
Alpine meadowtundra
Dicrostonyx torquatus
T
5
HudsonianCanadian (borealmontane)
Aplodontia rufa
Castor canadensis
Gulo gulo
Lepus americanus
Lepus townsendii
Marmota flaviventris
Martes americana
Microtus longicaudus
Microtus montanus
Microtus pennsylvanicus
Mustela erminea
Mustela nivalis
Myodes gapperi
Neotoma cinerea
Neovison vison
Ochotona princeps
Ondatra zibethicus
Phenacomys intermedius
Sorex cinereus
Sorex palustris
Spermophilus lateralis
Sylvilagus nuttallii
Vulpes vulpes
FU
R
F
FU
G
XM
F
MU
M
M
ER
M
F
X
RU
XM
R
MEU
F
RU
E
O
OE
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
2
4
5
4
6
14
3
2
7
2
5
2
2
15
5
9
2
4
2
4
7
5
9
Sonoran
Bassariscus astutus
Brachylagus idahoensis
Lemmiscus curtatus
Lepus californicus
Microtis californicus
Neotoma lepida
Perognathus parvus
Spermophilus mollis
(¼ townsendii)
Sylvilagus audubonii
Tamias minimus
Urocyon cinereoargenteus
FX
S
SG
O
GR
O
S
S
OU
F
F
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
2
4
7
5
2
4
2
5
þ
þ
þ
þ
þ
þ
þ
2
3
2
þ
þ
2
þ
þ
þ
Specialized niches
Wetriparian
Procyon lotor
Steppeprairie
Microtus ochrogaster
Mustela nigripes
Spermophilus richardsonii
Spermophilus
tridecemlineatus
Vulpes velox
R
G
G
G
G
þ
þ
þ
þ
þ
G
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
3
6
6
2
þ
6
Other
Mephitis mephitis
Mustela frenata
E
OR
þ
þ
þ
þ
þ
þ
7
9
92
Vol. 90, No. 1
JOURNAL OF MAMMALOGY
APPENDIX I.—Continued.
Wyomingb
Taxa
Spermophilus elegans
Thomomys bottae
Thomomys talpoides
Total taxa/site
a
Coloradoc
Habitata NT LBE BC LCC MC
þ
M
M
M
11
26
CR
Idahod
Nevadae
Californiaf
WB JC MSCT SCC SNC HCP HCH BRC
PC
SWL
Total
þ
þ
þ
22
11
3
14
þ
þ
þ
þ
13
21
29
16
11
þ
þ
5
9
2
3
6
þ
3
14
7
E ¼ edge; F ¼ forest; G ¼ grasslands; M ¼ meadow; O ¼ open; R ¼ riparian; S ¼ sagebrush; T ¼ tundra; U ¼ dense understory; X ¼ rocky.
NT ¼ Natural Trap Cave; LBE ¼ Little Box Elder Cave; BC ¼ Bell Cave; LCC ¼ Little Canyon Creek Cave; MC ¼ Mummy Cave.
c
CR ¼ Chimney Rock.
d
WB ¼ Wilson Butte Cave; JC ¼ Jaguar Cave; MSCT ¼ Moonshiner Carnivore Trap.
e
SCC ¼ Smith Creek Cave; SNC ¼ Snake Creek Burial Cave; HCP ¼ Hidden Cave Pleistocene strata, HCH ¼ Hidden Cave Holocene strata; BRC ¼ Bronco Charlie Cave.
f
PC ¼ Potter Cave; SWL ¼ Samwel Cave.
b