Chapter 6
NATIVE A MERICANS,
ECOSYSTEM
DEVELOPMENT, AND
HISTORICAL RANGE
OF VARIATION
Gregory J. Nowacki,1 Douglas W. MacCleery,2,3 and
Frank K. Lake4
1
USDA Forest Service, Eastern Regional Ofﬁce, Milwaukee, WI, USA
USDA Forest Service, Washington Ofﬁce, Forest Management (retired), Washington, DC,
USA
3
Alexandria, VA, USA
4
USDA Forest Service, Paciﬁc Southwest Research Station, Redding, CA, USA
2
Historical Environmental Variation in Conservation and Natural Resource Management, First Edition. Edited by John A. Wiens,
Gregory D. Hayward, Hugh D. Safford, and Catherine M. Giffen.
© 2012 John Wiley & Sons, Ltd. Published 2012 by John Wiley & Sons, Ltd.
76
Native Americans, ecosystem development, and historical range of variation
6.1
INTRODUCTION
In North America, human beings have not always
been considered “legitimate players” in natural ecosystems, often being dismissed as irrelevant, unnatural, or
simply “noise” in the system. This is in contrast to the
continents outside the Americas, where human history
is very long and disentangling human history from
ecosystem history is essentially impossible, not to
mention nonsensical (Bakker et al. 2000; Jackson &
Hobbs 2009). The debate over whether humans are
natural parts of North American ecosystems lingers
today (Comer 1997; Haila 1997; Hunter 1997), but in
our view, it is imperative to include humans in discussions of the ecological past, recognizing that human
inﬂuences on ecological history vary from signiﬁcant
to slight depending on the location and time frame
(Vale 2002). In this chapter, we describe the relative
importance of human effects on past ecosystems in
North America and, in light of the evidence, discuss
the relevance of human history to the historical range
of variation (HRV) concept. A spectrum of human–
environmental relations is discussed, including human
predation on wildlife populations, and the effects of
deliberate burning and agriculture/horticulture on
vegetation. Human–environmental relations vary
among ecosystems and cultures and are illustrated for
a representative set of ecoregions across the United
States.
6.2 HUMAN ENTRY INTO THE
“NEW WORLD ”
Throughout the Pleistocene, North American ecosystems reorganized rhythmically with glacial cycles
roughly every 100 000 years, driven by an array of
processes absent of humans (Pielou 1991; Delcourt
2002). The mix of drivers changed at the end of the
last glacial period when human arrival ushered in the
Holocene Epoch. Most Native Americans are the direct
descendants of northeast Asians (Meltzer 2003) who
entered the New World via the Bering Land Bridge
approximately 17–22 000 years ago (Mann & Hamilton 1995). Upon arrival, humans brought with them
three key things that would profoundly change the
American landscape: (1) tools (including weapons), (2)
dogs (Fiedel 2005), and (3) ﬁre (Stewart 2002).
Populations pooled in Eastern Beringia (Alaska and
Yukon Territory), a cul-de-sac of sorts, until ice sheet
77
retreat sometime between 13 000–14 000 years ago
(Mann & Hamilton 1995; Meltzer 2003). Once south
of the glacial barriers, migrants dispersed swiftly
across America, most likely in waves of genetically and
linguistically divergent tribal groups (Rogers et al.
1990). For instance, one of the early cultures, the
Clovis, may have expanded across North America in as
little as 200 years (Waters and Stafford 2007). The
Clovis peoples had the traits of big-game hunters, coupling a diverse toolkit, including ﬁre and large-ﬂuted
projectiles, with high mobility and broad habitat use.
Megafauna (large animals like mammoths [Mammuthus], mastodons [Mammut], and ground slouths
[Megatherium and Eremotherium]) were preferentially
sought as food. As conspicuous targets on the landscape that left abundant signs of their presence (trails,
dung, and broken branches), megafauna could be
pursued with relative ease, even as their numbers
dwindled (Haynes 2002). Aided with dogs, the ability
to track, engage, and slay megafauna and transport
meat from kill sites would have made Clovis groups
incredibly efﬁcient and lethal hunters (Fiedel 2005).
Because of all these factors, the Clovis cultures have
been implicated in Late Pleistocene megafaunal extinctions (Haynes 2002).
Megafaunal extinction at the hands
of humans
The extinction of megafauna was an exceptional ecological event with consequences still reverberating
through ecosystems today. A total of 34 megafaunal
genera went extinct in North America in the Late Pleistocene, including all mammals larger than 1000 kg
(Koch & Barnosky 2006). Numerous hypotheses have
been advanced to explain megafaunal extinction in
North America (Koch & Barnosky 2006). One of the
more durable explanations was offered by Martin
(1967), entitled the “Pleistocene overkill” theory. Since
initially proposed, this human-based extinction
hypothesis has endured scientiﬁc debate with ongoing
research and changing philosophies (Grayson &
Meltzer 2003; Fiedel & Haynes 2004). Skeptics claim
that severe climatic reversals and associated vegetation
shifts during the Pleistocene-Holocene transition were
to blame. But then, why were megafauna able to
survive numerous prior glacial-to-interglacial transitions only to succumb in the last one? The only apparent difference was human presence. Human-based
78
Issues and challenges
extinctions are not unprecedented, having occurred
across the globe over millennia (Martin & Steadman
1999; Koch & Barnosky 2006).
Extinction across a range of animals, from grazers
to browsers and generalists to specialists, strongly
argues against climate-based vegetation shifts as the
probable cause, but is wholly consistent with humanhunting models (Owen-Smith 1987; Koch & Barnosky
2006). Humans affected megafauna directly (hunting)
and indirectly (through competition and habitat alteration) in the midst of postglacial climate change.
Changing climate and habitat conditions may have
added stress to megafauna populations, but it was
humans who ultimately did them in (Haynes 2002).
Without human predation, it is unlikely that a Late
Pleistocene mass extinction would have occurred
(Koch & Barnosky 2006).
6.3 ECOSYSTEMS AFTER
MEGAFAUNA – HUMANS BEGIN TO
TAKE CONTROL
Megaherbivores were the keystone species of their time,
disproportionally affecting their environs and embedded food webs. Their ability to radically transform landscapes, open forest and shrub canopies, and promote
habitat diversity inﬂuenced a multitude of animals at
all trophic levels (Owen-Smith 1987). Large proboscideans (mammoths and mastodons) were particularly
adept at habitat modiﬁcation through grazing, browsing, trampling, and wallowing. Megaherbivore extinctions rippled through the animal kingdom, causing
further population reductions, food web collapses, and
extinctions among animals dependent on megafaunal
activities or presence. Openland-dependent, nonmigrating animals and large carnivores and scavengers
were most vulnerable (Haynes 2002).
As Johnson (2009, p. 2509) aptly states, “big herbivores have big effects on plants” and thus on community composition, structure, and patterns. Some
North American trees such as honey locust (Gleditsia
triacanthos), hawthorns (Crataegus), mesquite (Prosopis), and Osage-orange (Maclura pomifera) still carry
obsolete megaherbivore defenses in the form of stout
thorns (Johnson 2009; Bronaugh 2010). Upon extinction, many megaherbivore-limited food sources suddenly became plentiful (Janzen & Martin 1982); the
liberation of preferentially browsed trees, especially
hardwoods, is clearly seen in the pollen record (Gill
et al. 2009). Conversely, anachronistic species reliant
on megaherbivores for seed scariﬁcation and dispersal
declined, including Kentucky coffee tree (Gymnocladus
dioicus), honey locust, Osage-orange, pawpaw (Asimina
triloba), and persimmon (Diospyros virginiana) (Janzen
& Martin 1982; Barlow 2001). In the post-megafauna
world, the trajectories of ecosystem development were
markedly different than any preceding interglacial
period (Johnson 2009; Bronaugh 2010), leading to a
complete reorganization of vegetation with new species
assemblages and food webs. From this standpoint
alone, humans have profoundly affected ecosystem
development across the Americas since the onset of the
Holocene.
Humans and ﬁre
What characteristic best differentiates humans from
other species: language, intelligence, use of tools?
Some would argue it is the use of ﬁre, as no other
species has been able to master it (Goudsblom 1987).
Fire is an important evolutionary force – a force that
humans have used to shape environments across the
globe (Sauer 1975; Bond & Keeley 2005). The domestication of ﬁre was intrinsic to the development and
spread of human culture, dating back at least 100 000
years and possibly over 1 million years (Bowman et al.
2009; Pausas & Keeley 2009). Invariably during this
period, where there was man, there was ﬁre (Sauer
1975; Pausas & Keeley 2009). The people traversing
the cold Arctic environments surely brought ﬁre technologies to North America.
The spread of ﬁre- and weapon-toting humans
across North America left an indelible mark on the
paleoecological record. Disturbance regimes changed
from megaherbivory to human-based ﬁre. The rise of
ﬁre (as measured in charcoal) commensurate with
megafauna decrease (measured in dung fungal spores)
is reported throughout North America (Davis & Shafer
2006; Gill et al. 2009). An increase in ﬁre may have
compensated for ecological changes expected from
declines in herbivory, as ﬁre can be considered an “herbivore” of sorts (see Bond & Keeley 2005). The switch
from megaherbivory to a human-mediated ﬁre regime
led to a new set of plant communities, resulting from
the rebound of plants formerly suppressed by herbivory (relaxation), reduction of anachronistic plants,
and the promotion of ﬁre-adapted plants (Gill et al.
2009; Johnson 2009).
Native Americans, ecosystem development, and historical range of variation
The rise of agriculture
An extensive switch in human diet ensued after megafaunal extinction. Human population in North
America likely declined responding to the combined
loss of a signiﬁcant food source (megafauna) and deterioration of climate during the Younger Dryas. After
megafaunal extinction, people switched to smaller
Folsom points designed for smaller prey. Subsistence
technologies diversiﬁed, allowing for greater utilization
of resources, especially plants.
With the advent of agriculture about 5000 BP,
eastern North America became one the ﬁve major
centers of independent agricultural development in
the Americas (Smith & Yarnell 2009). During this
time, people transitioned from highly mobile huntergatherers to more sedentary, agriculture-based societies (at least in those areas that supported agriculture).
By 3800 BP, agriculture in the eastern United States
had become more intense, sophisticated, and responsible for an increasing share of total human nutrition.
Human societies responded by growing and becoming
more complex. Agrarian lifestyles ﬂourished in warmer,
more productive southern portions of the continent.
Especially after 800 AD, Three Sister agriculture (composed of maize [Zea], beans [Phaseolus], and squash
[Cucurbita]) arose in southeastern North America,
eventually spreading to many parts of what is now the
United States by the sixteenth century.
Across all cultural phases of the Holocene, ﬁre continued to be a key tool in food acquisition, especially in
maintaining open habitats, fostering berry- and nutproducing shrubs and trees, game hunting, and land
clearance for gardening and habitation (Doolittle
2000; Williams 2000; Abrams & Nowacki 2008).
Upon their arrival along the coasts, Europeans did not
“discover” an untrammeled wilderness but rather a
kaleidoscope of ecosystems reﬂecting the many native
peoples who lived there (Heizer 1978; Patterson & Sassaman 1988; Suttles 1990). Europeans found the preconditioned landscapes (old Indian villages, gardens,
and ﬁre-maintained clearings) most suitable for settlement as they afforded the best chances for survival
(Mann 2005). Such locations were often the most productive and geographically strategic on the landscape.
Butzer (1990, p. 27) wrote that in North America,
there existed “a pre-European cultural landscape,
one that represented the trial and error as well as
the achievement of countless human generations. It
is upon this imprint that the more familiar Euro-
79
American landscape was grafted, rather than created
anew.”
6.4 HUMAN-MODIFIED LANDSCAPES
AT THE EVE OF EUROPEAN CONTACT
By the time Europeans arrived in America, indigenous
peoples had occupied the continent for many millennia. Human effects on North American landscapes
were pervasive in the most productive regions (Southeast, West Coast) and limited in the least productive
regions such as the boreal North and interior West
(Pyne 1982; Patterson & Sassaman 1988; Vale 2002).
Pre-European population density would appear to be a
good proxy for human ecological impact. Tribal cultures were an integral part of productive landscapes,
serving as active change agents who intentionally
managed resources (Stewart 1963; Nicholas 1988).
Native Americans affected ecosystem development in
enumerable ways, including hunting, gathering,
ﬁshing, agriculture, arboriculture (active planting and
dissemination of desired woody species), wood gathering, village and trail construction, and habitat manipulation (Williams 2000; Abrams & Nowacki 2008). Fire
was used for many of the aforementioned activities,
ultimately being the tool of choice to manage landscapes for many socioeconomic beneﬁts (Pyne 1983;
Williams 2000).
Through examples spanning North America from
east to west, the remainder of this chapter illustrates
the diverse and often profound inﬂuence Native Americans had on ecosystems across the continent. We geographically organize our review of historical human
effects on ecosystems, using standard ecological divisions of North America as a template (Cleland et al.
2007; Fig. 6.1).
Eastern North America
A large percentage of global vegetation does not reﬂect
climax conditions set by climate, but rather represents
subclimax conditions largely driven by ﬁre (Bond et al.
2005). This was certainly the case in the eastern
United States where ﬁre-dependent communities, such
as tallgrass prairies and oak (Quercus), pine (Pinus),
and oak-pine woodlands, dominated at the time of
European contact (Frost 1993; Hamel & Buckner
1998; Nowacki & Abrams 2008). These principally
80
Issues and challenges
Marine
Mediterranean
Warm Continental
Temperate Desert
Hot Continental
Temperate Steppe
Subtropical
Tropical/Subtropiacl Desert
Savannah
Tropical/Subtropiacl Steppe
Prairie
N
0
250
500
1000 km
Fig. 6.1 Ecological Divisions of the continental United States (source: Cleland et al. 2007).
open vegetation types sharply contrast with the closedcanopy broadleaf forests that would have otherwise
prevailed (Bond et al. 2005). Fire was the principal
mechanism controlling vegetation expression in the
East, as without it most forested systems would have
been composed of late-successional species such as
hemlock (Tsuga), ﬁr (Abies), maple (Acer), basswood
(Tilia), beech (Fagus), and magnolia (Magnolia). Fires
were largely attributed to Native Americans as most
were dormant season burns when lightning was at a
minimum (Barden 1997; Lorimer 2001; Abrams &
Nowacki 2008). Moreover, once started, Native Americans had little incentive or means to put ﬁres out,
hence explaining the large spatial extent of ﬁredependent ecosystems across the East.
At earliest European contact, human modiﬁcations
of landscapes were not uniform along the East Coast,
but followed a north-south productivity and human
population gradient (Driver & Massey 1957; Patterson
& Sassaman 1988). In the far north, Native Americans
were more oriented to hunting, ﬁshing, and gathering
since the cool, short growing season was less conducive to agriculture. The low carrying capacity of the
land restricted human populations, and in turn their
environmental effects. One exception was wild rice
(Zizania) husbandry, which thrived within this lake-
Native Americans, ecosystem development, and historical range of variation
studded region. There is some evidence that maize consumption and possible cultivation took place in the
southern portions of the boreal forest, but again environmental effects would have been minimal (Boyd &
Surette 2010). Human ignitions only supplemented
the principally lightning-based ﬁre regime.
Warm Continental Division: conifer-northern
hardwood systems
Land impacts were more conspicuous where boreal
conifers intermingled with temperate broadleaf trees
(Warm Continental Division, Fig. 6.1), although effects
were primarily conﬁned to areas around villages,
encampments, and trails (Campbell & Campbell 1994;
Clark & Royall 1995). Here, the prevailing cool and
moist climate coupled with the pyrophobic hardwoods
(wet, ﬂaccid leaf litter) greatly suppressed ﬁre, thwarting human’s most potent land-altering tool. Consequently, Native Americans focused on smaller, more
manageable burns immediately surrounding their
river- and lake-side villages, creating disturbance
patches or cultural “islands” within a sea of climax
northern hardwoods (Table 6.1; Mann et al. 1994).
Western New York and northwestern Pennsylvania
are representative, where small clearings and oakdominated forests occurred in areas previously occupied by the Iroquois, especially along rivers and
pathways (Ruffner & Abrams 2002; Black et al. 2006).
Here, the small and localized effects of Native Americans were reﬂected in land surveys, with activities
recorded on <1% of the bounds surveyed in the 1790s;
since this land survey followed Iroquois depopulation,
this estimate is biased low but remains an indication of
the relatively small human footprint (Marks & Gardescu 1992). The amount of land disturbance undoubtedly oscillated over time contingent with Native
American movements and population levels (Ruffner &
Abrams 2002; Stambaugh & Guyette 2006).
The intensity of human impacts radiated from cultural centers in concentric rings, with ﬁelds within
1 km, fuelwood collection and burning for berry production within 4–5 km, and foraging and small game
hunting within 6–8 km (Williams 1989; Black et al.
2006). One string of villages along the shores of Green
Bay (Wisconsin) is illustrative of the human footprint
on the northern hardwood matrix (Dorney & Dorney
1989). Here, Potawatomi and Winnebago peoples
established an agricultural-horticultural landscape of
81
ﬁelds, open lands, and oak savannas and forests.
Burning apparently created these vegetation conditions as soils, topography, or climate did not appreciably differ between the human-modiﬁed locale and the
distal northern hardwoods.
Similar impacts were found in southern Ontario
(Clark & Royall 1995; Munoz & Gajewski 2010), where
the Native American footprint probably did not exceed
3.2% of the total land base (Campbell & Campbell
1994). At Crawford Lake, sediment analysis showed
that increases in charcoal corresponded with increases
of ﬁre-dependent pine (Pinus) and oak (Quercus), the
appearance of maize, weedy grasses, and purslane
(Portulaca) (distinct cultural markers), and decreases of
maple and beech from ca. 1350 to 1650 AD. Burning
was cited as the primary factor in converting northern
hardwoods to open systems of pine and oak. Furthermore, human ignitions have been implicated in the
extension of oak-hickory forests up major river valleys
along the southern margin of northern hardwood
system (Cogbill et al. 2002; Black et al. 2006). Acorns
were a highly prized, storable carbohydrate source that
supplemented Native American diets, motivating land
disturbances that promoted oak and associating
species (e.g. blueberries) within the northern hardwood complex (Abrams & Nowacki 2008). Dunham
(2009) found archeological sites disproportionally
associated with oak habitats in the eastern Upper
Peninsula of Michigan. Overall, human land alterations promoted biological and landscape diversity and
certainly increased the abundance of shade-intolerant,
ﬁre-adapted, and culturally important plants in these
Northwoods.
Hot Continental Division: central and
Appalachian hardwoods
Human land modiﬁcations ﬂourished south of the
cool northern hardwood system where much of the
area was either directly or indirectly under Native
American control. The favorable climate of the Hot
Continental Division (Fig. 6.1) supported agriculture
and acorn- and nut-based cultures. Catchment analysis revealed that local cutting and burning practices
(for agriculture, fuelwood, and construction materials)
increased the representation of hickory (Carya), walnut
(Juglans), and black locust (Robinia pseudoacacia) (Black
& Abrams 2001), whereas broadcast burns led to the
dominance of oak (Quercus), hickory, and American
82
Issues and challenges
Table 6.1 Probable climax vegetation versus actual dominant vegetation at the time of European settlement by
Ecological Division.
Division
Climax vegetation1
Dominant presettlement
vegetation2
Native American
Inﬂuence3
Warm Continental
Hot Continental
Conifer-northern
hardwoods
Mixed mesophytic forests
Conifer-northern
hardwoods
Oak and oak-pine
woodlands
Subtropical
Beech-magnolia forests
Southern pine woodlands
Prairie
Mixed mesophytic and
oak forests
Tallgrass prairie
Temperate Steppe
Shrublands
Short and mid-grass
prairie
Temperate Desert
Sagebrush-grass,
ponderosa pine, and
shade-tolerant conifers
Shrublands and pinyonjuniper
Sagebrush-grass and
ponderosa pine
Local; limited to settlements
and travel corridors
Widespread; landscape burns
through Native American
ignitions
Widespread; landscape burns
through Native American
ignitions
Ubiquitous; prairies wholly
dependent on Native
American burning
Widespread; Native American
ignitions increased ﬁre
frequency to help support
grasses
Local to widespread
(depending on terrain)
Tropical/
Subtropical
Desert/Steppe
Marine
Mediterranean
Grasslands and pinyonjuniper
Western hemlockDouglas-ﬁr forests
Douglas-ﬁr forests
Mixed conifer-oak forests
Oak woodlands and
chaparral
Local; effects concentrated
on the most productive
areas
Local; prairies and shrublands
maintained by Native
American burning.
Widespread; open
communities maintained by
Native American burning
1
Postulated climax vegetation (uplands) in accordance with the disturbance regime without humans.
Actual dominant vegetation (uplands) in accordance with the disturbance regime with humans.
3
Stewart (2002), Abrams and Nowacki (2008), Williams (2000), and references therein.
2
chestnut (Castanea) over much of the surrounding
landscape (Delcourt & Delcourt 1997; Nowacki &
Abrams 2008). Without human-based ﬁre, much of
the region would have been dominated by shadetolerant mesophytic trees, such as maple, beech, and
basswood (Lorimer 1992; Table 6.1). Interestingly, the
interface between oak-dominated ecosystems of this
division and the northern hardwood ecosystems to the
north was probably more of a function of anthropogenic ﬁre than climate (Cogbill et al. 2002).
Descriptions of the open character of this region
were very common in the notes of early observers.
John Smith commented that around Jamestown, Virginia “a man may gallop a horse amongst these woods
any waie, but where the creekes and Rivers shall
hinder” (Williams 1989, p. 44). Andrew White, on an
expedition along the Potomac in 1633, observed that
the forest was “not choked with an undergrowth of
brambles and bushes, but as if laid out in by hand in a
manner so open, that you might freely drive a four
horse chariot in the midst of the trees” (Williams
1989, p. 44). Such observations of the open nature of
oak forests from the East Coast through the Ozarks are
typical of those of most early observers, who commonly spoke of the ease of riding a horse or driving a
wagon under a park-like canopy.
A mix of grasslands and open woodlands was maintained speciﬁcally for game, an important protein
source for indigenous peoples (Abrams & Nowacki
2008). There is evidence that Native Americans were
Native Americans, ecosystem development, and historical range of variation
encouraging the eastward migration of bison (Bison
bison) through habitat manipulation at the time of
European contact (Rostlund 1960; Pyne 1982). Sizable
grasslands surrounded by oak-dominated woodlands
once existed here. In Kentucky, a vast grassland on the
Pennyroyal Plateau measured approximately 249 km
(155 m) long and 19 km (12 m) wide (Lorimer 2001).
In Virginia, the Shenandoah Valley was one vast grass
prairie covering more than 2590 sq km (1000 sq mi)
where Native Americans burned annually (Van Lear &
Waldrop 1989). After burning by indigenous peoples
ceased, much of this area reverted to forest and the
early white settlers had to clear land that had only
recently been prairie (Rostlund 1957). R.C. Anderson
(1990, p. 14) writes that the eastern prairies and
grasslands “would mostly have disappeared if it had
not been for the nearly annual burning of these grasslands by the North American Indians.” Indeed, the
existence of the Prairie Peninsula (Prairie Division; Fig.
6.1) in the humid East is thought to be largely an artifact of Native American burning (Anderson 2006).
Where not plowed or pastured by Europeans, the open
character of the land quickly reverted to dense, closedcanopy oak forests due to ﬁre suppression. Under
today’s subdued ﬁre regime, these oak forests are
undergoing mesophication and converting to climax
forests of shade-tolerant maple, beech, and elm (Ulmus)
with little understory diversity (Table 6.1) (Nowacki &
Abrams 2008).
Subtropical Division: southern pines and
associated ecosystems
At the time of European contact, the ecological effects
of indigenous peoples were likely more signiﬁcant in
the Southeast than anywhere else in North America.
This was due to relatively high population densities
(Williams 1989), the widespread application of agriculture, as well as plant communities and climatic conditions that facilitated human use of ﬁre.
It is difﬁcult to accurately determine ecological conditions at European contact (1500 AD) as there were
few European observers in this region at that time. One
of the most important was the Hernando de Soto expedition. From 1539 to 1542, de Soto (with his 600 men,
200 horses and 300 swine) pillaged, plundered, and
inadvertently spread European diseases from Florida to
North Carolina, west across the Mississippi River, then
down to the Gulf of Mexico (Thomas et al. 1993; Mann
83
2005). Even with its large numbers of men and
animals, the de Soto expedition moved with relative
ease throughout the Southeastern landscape, and
chroniclers wrote of expansive agricultural ﬁelds, open
park-like forests, numerous villages, and large numbers
of people (Swanton 1939; Rostlund 1957).
In what is now Alabama, a de Soto chronicler
reported that the land was “so fertile and thickly populated that on some days the Spaniards passed 10 or 12
towns, not counting those that lay on one side or the
other of the road” (Rostlund 1957, p. 385). Upon
arriving at the Mississippi River, de Soto found a landscape teeming with humans. The river itself was lined
with villages (Mann 2005). Eerily, by the time the next
explorer passed through this area more than a century
later (La Salle in 1682), the entire valley had been radically transformed into a relatively silent place (Mann
2005). Where de Soto had observed scores of villages,
expansive agricultural ﬁelds, and high human populations, La Salle found mostly forest with very few people
or villages. The country had been depopulated (80–
95%) by European diseases, and the ecology of the area
substantially altered (Young & Hoffman 1993; Mann
2005; Scharf 2010).
Landscapes cleared for agriculture or routinely
burned had two or more centuries to recover before the
ﬁrst waves of permanent Euro-American settlers
arrived to ﬁnd landscapes that were more “pristine”
than they had been in more than a thousand years
(Denevan 1992; Scharf 2010). In addition to the rapid
succession of extensive old ﬁelds, the depopulation of
indigenous peoples undoubtedly led to changes in wildlife populations as a major predation source was substantially reduced. For instance, bison expanded as far
south as Florida and as far east as Virginia and Pennsylvania; the large numbers of bison transformed
many areas as they grazed, creating large wallows and
well-worn migration corridors, some of which remain
visible today (Rostlund 1960; Belue 1996).
Eyewitness reports describing ecological conditions
in the Southeast become more common after 1600
(Smith 1616; Lindestrom 1656, Lederer 1672, Catesby
1731; Bartram 1791). Many of these writers noted
extensive “ancient” Indian plantations and abandoned
ﬁelds extending for kilometers along rivers (see in particular Bartram 1791). The most common ecological
conditions reported by these observers were open
forests, interspersed with grasslands and meadows,
with extensive cane lands along the rivers. Bartram’s
journals contain numerous references to “delightful
84
Issues and challenges
groves” of open grown “stately forests” of oak, ash,
hickory, walnut, and so on, as well as “vast open
forests” (Bartram 1791). References to dense forests of
late successional species are rare indeed. Rostlund
(1957, p. 408) concludes that “the open, parklike
appearance of the woodlands, undoubtedly the most
common type of forest in the ancient Southeast, was
mostly the work of man.”
Frequent forest burning did more than create open
stands of shade-intolerant trees, but created grasslands where forests would have otherwise existed.
There are many references to treeless areas in the early
literature, which were often referred to as “barrens,”
“plains,” “meadows,” or “savannahs.” In Bartram’s
journal, there are numerous separate references to
“vast meadows,” “extensive savannas,” and “large
grassy plains,” some of which were reported to be
many kilometers in length (Rostlund 1957). Bartram
reported that the Alachua Savanna in northern Florida
was “a level green plain, above 15 m (24 km) over, 50 m
(80 km) in circumference, with scarcely a tree to be
seen” (Rostlund 1957, p. 408).
Canebrakes were also a major feature of Southern
bottomlands at the time of European settlement (Platt
& Brantley 1997). Indigenous people valued cane for
food, shelter, baskets, and tools, especially weapons
(Hamel & Chiltoskey 1975). William Bartram repeatedly remarked on canebrakes during his southeastern
travels, describing “vast cane meadows,” “widespread
cane swamps,” or “an endless wilderness of canes”
(Platt & Brantley 1997, p. 10). The area of canebrakes
declined rapidly in the eighteenth century from altered
disturbance regimes by Euro-Americans, including
cattle grazing, agricultural displacement, and changes
in ﬁre frequency. Today, cane has been virtually eliminated from the Southeastern landscape and is a contributing factor in the extinction of Bachman’s warbler
(Vermivora bachmanii) (Remsen 1986).
At the time of European contact, the Coastal Plain
was dominated by open stands of large pines (Bartram
1791). Within the range of the ﬁre-tolerant longleaf
pine, which covered about 37 million hectares, ﬁresensitive southern mixed broadleaved forests (beech,
magnolia, semi-evergreen oaks) were restricted to
moist, ﬁre-protected locations (Frost 1993). In the
southern portion of longleaf pine distribution, summer
lightning ﬁres were sufﬁcient to maintain longleaf pine
(Komarek 1964), but burning by indigenous people
likely contributed to range extension of longleaf and
other southern pines into topographically dissected
areas where it would not otherwise have occurred
(Frost 1993). By the early 1990s, Frost (1993) estimated that about 1 million hectares remained in naturally regenerated longleaf pine, with only about
270 000 ha (<0.7% of the original range) in a condition similar to the classic open-grown, ﬁre-maintained,
longleaf pine-wiregrass community. Like canebrakes,
the longleaf pine/wiregrass community is currently
considered a critically endangered ecosystem (Noss
et al. 1995).
The Great Plains and interior west
Native American populations became progressively
more sparse west of the Mississippi River across the
Great Plains (Temperate Steppe Division) through the
Rocky Mountains (Temperate Desert Division; Fig.
6.1), essentially following an aridity gradient. Together
with changes in topography, Native American–
environmental relations varied broadly across this
western expanse. The relatively ﬂat and seasonally dry
conditions in the center of the continent were naturally conducive to extensive ﬁre. The vast grasslands
that epitomized the Great Plains were promulgated by
frequent surface burning (Wright & Bailey 1980), and
fed large ungulates such as bison, elk (Cervus canadensis), deer (Odocoileus), and later horses (Equus). Ungulate populations were regulated by hunting where
denser Native American populations existed, perhaps
to the point of being largely relegated to the outskirts
of tribal territories (Martin & Szuter 1999; Kay 2007).
Well versed in prey–habitat relations, indigenous
peoples promoted preferred habitat conditions through
burning and were the principal igniters outside the
thunderstorm season (April to September) when
optimal fuel conditions (dry grass) existed. Without the
addition of human ignitions, the Great Plains may
have been substantially shrubbier, supporting fewer
large herbivores (Arno & Gruell 1983; Stewart 2002;
Table 6.1). Post-European agriculture, grazing, and
ﬁre suppression adversely affected the species composition and diversity of the Great Plains.
In the Rocky Mountains (Temperate Desert Division),
resources important to indigenous peoples were geographically spread from river valley to alpine meadow.
Evidence suggests that management of resource
patches by indigenous people was common, although
at decreasing intensity with elevation (Barrett & Arno
1982; Hessburg & Agee 2003). Many tribes practiced
Native Americans, ecosystem development, and historical range of variation
seasonal rounds, migrating to higher elevations during
the summer, and back to winter villages in the lowlands. Along the way, both the ﬂora and fauna of these
mountainous ecosystems were affected by Native
American hunting, plant gathering, and burning. Early
observations of western ﬁres compiled by Gruell (1985)
revealed that many were set by indigenous peoples,
especially at lower and middle elevations. Here, intentional burning helped create and maintain a mosaic of
stand conditions across the landscape. Periodic burning
helped promote root and berry crops, materials for basketry, forage for prey animals, and improved visibility
for hunting (Stewart 2002). However, given the inherent severity of mountain and high desert environments
(rugged topography, short growing seasons, temperature extremes, snow, and aridity), Native Americans
probably had a much lighter touch in this region relative to other, more productive areas of North America
(Vale 2002). Nevertheless, human ignitions were
embedded in historical ﬁre regimes that supported a
diverse vegetation mosaic – a mosaic now being
unraveled by ﬁre suppression (Pyne 1982; Arno &
Gruell 1983; Brown & Hull-Sieg 1996). Repeat photography clearly shows the dramatic recent increases in
forest cover in many western landscapes (Progulske
1974; Gruell 1983; Skovlin & Thomas 1995).
West Coast
Marine Division: temperate rainforests
Lightning has been accepted as the main ignition source
of ﬁres in this topographically diverse region, with frequency of strikes increasing with distance from coast
and with elevation (Agee 1993; van Wagtendonk &
Cayan 2008). Fires of the mesic forests in Coast Ranges
and Cascades were infrequent, stand-replacing, highseverity events (Agee 1993; Lertzman et al. 2002),
although mixed severity and more frequent lowmoderate severity ﬁre regimes existed in mixed coniferhardwood vegetation and grassland communities (Agee
1993; Weisberg & Swanson 2003; Taylor et al. 2008).
Where summer lightning ﬁres were not adequate to
foster and maintain the desired condition, tribal burning
was used. The extent to which Native American ﬁres
affected historical ﬁre regimes of these mesic coniferdominated forests is debated (Agee 1993; Boyd 1999).
The longevity and extent of burning practices by
diverse tribal groups across the Paciﬁc Northwest and
85
California is not clear from archeological interpretations (Ames & Maschner 1999; Boyd 1999). Late
Holocene tribal groups did not practice sedentary agriculture, but rather a proto-agriculture/horticulture
integrated with ﬁre regimes (Lewis 1993; Boyd 1999).
Vegetation nearest villages, trail systems, and remote
resource collection areas (i.e. camps) was selectively
modiﬁed by altering the frequency, seasonality, and
extent of ﬁres (Boyd 1999). Cultural practices sought
to increase wildlife forage, root, seed, nut, and berry
crops, enhance basketry materials, and improve
hunting opportunities (Boyd 1999).
Mediterranean Division: oak balanocultures
Across the major mountain ranges and valleys of California, the establishment and increased abundance of
oaks (Quercus and Lithocarpus) took place in the MidHolocene approximately 9000–10 000 years ago (West
et al. 2007; Briles et al. 2008). Oaks likely reached
their present distribution during the Late Holocene
(West et al. 2007).
Frequent lower intensity ﬁres promoted oakdominated habitats. Dendroecological studies indicate
that ﬁres occurred primarily during the summer seasonal drought, typical of the lightning ignitions (van
Wagtendonk & Fites-Kaufman 2006). However, there
is also evidence of spring burns, which suggests human
ignitions (Skinner et al. 2009).
Estimates of precontact (circa 200 years BP) ﬁrereturn intervals range from annual to decadal events
for Oregon white oak/grassland communities (McDadi
& Hebda 2008), and from 5 to 20 years for mixed oak/
conifer-hardwood forests of California (Skinner &
Taylor 2006). Charcoal and pollen analyses suggest a
reduction in ﬁre frequency during European settlement
(McDadi & Hebda 2008), leading to the conversion of
oak-dominated savannas to closed-canopy forests of
more mesic, ﬁre-sensitive species. This change occurred
despite broad climatic similarity between pre- and postEuropean eras (Byrne et al. 1991).
The rapid change in grassland/oak vegetation coincident with European settlement during a relatively
stable climate provides a striking contrast to the persistence of grassland/oak vegetation over long periods
of variable climate during Native American management. Subsistence economies (DeLancey & Golla 1997)
associated with oak grasslands remained fairly constant over the last 2500 years in the Sacramento and
86
Issues and challenges
San Joaquin valleys (Elasser 1978). Those tribal groups
inhabiting inland oak/grass-dominated valleys are
thought to have been deliberately broadcast-burning
vegetation by 3000, if not 5000 years BP (Weiser &
Lepofsky 2009). Archeological evidence indicates the
majority of nonmarine/littoral-adapted tribal groups
developed important associations with oak-dominated
habitats. Milling stones and related lithic tools (e.g.
mortars and pestles) used for processing seeds, nuts, or
other foods by tribal groups suggest an adaption to use
acorns as early as 7000 years ago in the California
North Coast Range and 4000 years BP in the San Francisco Bay Area, Sacramento Valley, and southern California (Elasser 1978; Keeley 2002; Arnold et al. 2004).
By 1300 years ago, acorns were a cultural mainstay
in the Sacramento/San Joaquin Valleys and in southern California (Elasser 1978; Arnold et al. 2004). The
bow and arrow was introduced by 1500–2000 years
ago, and coincides with other technological changes
(e.g. mortars and pestles) associated with diversiﬁed
subsistence economies (acorns, roots, seeds/nuts, ﬁsh,
game, etc.). The use of ﬁre by tribal groups to promote
oak dominance coincided with the need to counteract
natural successional tendencies, and the need to maintain tribal economies dependent on oaks and associated
species that provided valued food, materials, and medicines; various tribal groups exhibited proprietor claims
to tracts of oaks and associated resources (game and
bulbs/roots) within these habitats (Anderson 2007;
McDadi & Hebda 2008). Inland areas codominant with
oaks, and especially traversable ridge systems with
southern exposure, were kept open through burning
by tribal groups that depended on resources found in
early seral oak grasslands (Boyd 1999; Keeley 2002).
A combination of biophysical conditions and tribal
burning likely fostered oak dominance and persistence.
It is unknown just how much of the oak-dominated
landscape was inﬂuenced by tribal burning (Boyd 1999;
Anderson 2007). Oak habitats have greatly declined
since Euro-American settlement due to the cessation of
tribal burning, grazing of livestock, ﬁre suppression,
industrial forestry practices (favoring conifers), and
urbanization (Hosten et al. 2006; Anderson 2007).
6.5 NATIVE AMERICANS AND THE
HRV CONCEPT
Precontact Native American cultures were sophisticated, dynamic, diverse social organizations with indi-
viduals who knew how to alter their surroundings for
their beneﬁt (Pyne 1983). Since their postglacial
arrival, human populations and their scope of inﬂuence increased as they dispersed across the Americas,
reaching a peak at the time of European contact (Mann
2005; Scharf 2010). By that time, human population
density closely paralleled gradients in land productivity, ranging from low to high densities from north to
south in eastern North America and from mountains
and deserts (low) to fertile valleys and coastlines (high)
in the West. As such, the geographical inﬂuences of
Native Americans were not uniform, but reﬂected differences in populations, land use, cultural traditions,
available resources, climate, and vegetation ﬂammability (Kimmerer & Lake 2001; Stewart 2002; Vale 2002).
In North America, literature illustrating the various
roles of early humans in environments has not been
well integrated into our ecological knowledge. The
reasons are many, including historic and cultural
biases and political agendas, as well as the challenges
associated with obtaining clear scientiﬁc evidence of
environmental conditions 300 or more years ago
(Stewart 1963; Pyne 1982; Williams 2000). The
important role humans played was largely overlooked
by most early ecologists, who described North American landscape in terms of static “climax” systems consistent with the prevailing philosophy at the time. Early
paleoecologists also largely ignored the human factor,
tying Holocene species movements and ecosystem
reorganization exclusively to climate (Delcourt & Delcourt 2004; Scharf 2010). Shades of this oversight
continue today, through our intense focus on climate
change, which tends to assume that vegetation (baseline data for climate-change models) is solely reﬂective
of climate. Indeed, a new synthesis is needed integrating ecology, paleontology, and archeology to discern
the complex relations between ecosystems and embedded humans (a panarchical approach; Delcourt & Delcourt 2004). Fortunately, the acknowledgment of past
humans as an ecological factor is gaining momentum,
and is reﬂected in the use of “historical range of variation” rather than “natural range of variation.” In this
regard, humans are essentially acknowledged as the
change agents that put the “H” in the HRV concept.
The preceding sections chronicle a rich array of
Native American inﬂuences on ecosystems across time
and space, beginning with their continental arrival as
the new top predator. By employing weapons, dogs,
and efﬁcient hunting strategies, Native Americans
proved lethal against an unwary megafaunal prey
Native Americans, ecosystem development, and historical range of variation
base, and played a leading role in the rapid demise of
many large herbivore species. The removal of megaherbivores had a liberating effect on plant life (especially trees) in the midst of postglacial revegetation of
the North American continent. Native Americans also
represented a new ignition source – an ever-constant
source of ﬂame that was less constrained (spatially,
seasonally) than natural ignitions (Stewart 2002;
Abrams & Nowacki 2008). Fire ultimately became the
most powerful tool for landscape manipulation (Sauer
1975; Kimmerer & Lake 2001), which Native Americans used with purpose and facility. Later, as huntergatherer societies shifted toward more sedentary
lifestyles with growing technology, direct habitat
manipulation through agriculture and horticulture
took place where climate and soils allowed. Through
these three factors alone, Native American impacts
spanned spatiotemporal scales from continental
(megafaunal demise and subsequent trophic effects;
Early Holocene onward) to landscape (broadcast
burning; Mid-Holocene onward) to local (agriculture/
horticulture; Late Holocene).
Not all human-related ecological disturbances are
equally important to HRV analyses. Application of historical ecology generally relies on time-space scales
relevant to current land-management decisions, focusing on shorter, more recent periods with relatively
stable climates, vegetation, and disturbance regimes at
a particular locale. Coupled with inherent data limitations (e.g. a fading data record with time), timelines are
often restricted to the last thousand years or less. To
help integrate the consequences of Native American
land use into HRV analyses, data collection at a
minimum should identify and record principal landuse activities associated with former tribal territories.
Linking human-based disturbances (type, frequency,
intensity) with plant community structure and composition allows a more complete understanding of ecosystem dynamics, further enlightening the science
behind restoration.
Why is it important to understand the role of indigenous people in ecosystem dynamics described by HRV
analyses? Altered disturbance regimes affecting many
North American ecosystems may result in unsustainable conditions. The “pristine myth” is an enormous
cultural barrier to purposeful human intervention in
these systems (Hamel & Buckner 1998). Many wildernesses and protected “natural” areas exhibit vegetation
conditions at odds with what would have been expected
in pre-European landscapes and are on unprecedented
87
ecological trajectories. Without active human intervention, open woodlands of shade-intolerant species
(systems that have dominated many landscapes for
millennia) will continue to be replaced by closed forests
of shade-tolerant species (Nowacki & Abrams 2008).
Developments in historical ecology have improved
our understanding of past disturbance regimes, which
are vital to predicting vegetation succession and
dynamics. Indeed, many rare and endemic plant and
animal species in the United States are disturbance
dependent, and many of those historical disturbances
were associated with Native American activities. A
large proportion of the endangered ecosystems listed
by Noss et al. (1995) are ﬁre-dominated ecosystems.
These include prairies, pine savannahs and barrens,
tropical hardwood hammocks, and shrublands. Of 21
rare communities listed in the Ozark-Ouachita Highlands Assessment, the decline of nine was attributed to
ﬁre exclusion (USDA 1999). Nationwide, Owen and
Brown (2005) found that of 186 federally listed, proposed, and candidate plant species, 25% required ﬁre,
35% tolerated ﬁre, 38% were not affected by ﬁre, and
only 2% were adversely affected by ﬁre.
Resource managers are largely powerless to counter
the ecological effects of the substantial land-use
changes that have occurred in North America over the
last three centuries. But they are not powerless to recognize and address through management activities the
ecological effects of altered disturbance regimes. An
understanding of natural and human inﬂuences on
the development of historical landscapes is critical
to effectively planning and executing projects designed
to restore or conserve rare and endemic species and
ecosystems (Hamel & Buckner 1998; Scharf 2010).
Developing an appreciation of the past roles of humans
in modifying “natural” ecosystems in North America
is also a ﬁrst step toward better understanding and
managing the roles humans will play in future
ecosystems.
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