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Elephants (and extinct relatives) as earth-movers and ecosystem engineers


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Modern African elephants affect habitats and ecosystems in significant ways. They push over trees to feed on upper branches and often peel large sections of bark to eat. These destructive habits sometimes transform woody vegetation into grasslands. Systems of elephant trails may be used and re-used for centuries, and create incised features that extend for many kilometers on migration routes. Elephants, digging in search of water or mineral sediments, may remove several cubic meters of sediments in each excavation. Wallowing elephants may remove up to a cubic meter of pond sediments each time they visit water sources. Accumulations of elephant dung on frequented land surfaces may be over 2kg per square meter. Elephant trampling, digging, and dust-bathing may reverse stratigraphy at archeological localities. This paper summarizes these types of effects on biotic, geomorphic, and paleontological features in modern-day landscapes, and also describes several fossil sites that indicate extinct proboscideans had very similar effects, such as major sediment disturbances.
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Elephants (and extinct relatives) as earth-movers and ecosystem engineers
Gary Haynes
Department of Anthropology, University of Nevada-Reno, Reno, NV 89557-0096, United States
abstractarticle info
Article history:
Accepted 4 April 2011
Available online 23 June 2011
Ecosystem engineering
Landscape sculpting
Modern African elephants affect habitats and ecosystems in signicant ways. They push over trees to feed on
upper branches and often peel large sections of bark to eat. These destructive habits sometimes transform
woody vegetation into grasslands. Systems of elephant trails may be used and re-used for centuries, and
create incised features that extend for many kilometers on migration routes. Elephants, digging in search of
water or mineral sediments, may remove several cubic meters of sediments in each excavation. Wallowing
elephants may remove up to a cubic meter of pond sediments each time they visit water sources.
Accumulations of elephant dung on frequented land surfaces may be over 2 kg per square meter. Elephant
trampling, digging, and dust-bathing may reverse stratigraphy at archeological localities. This paper
summarizes these types of effects on biotic, geomorphic, and paleontological features in modern-day
landscapes, and also describes several fossil sites that indicate extinct proboscideans had very similar effects,
such as major sediment disturbances.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
Large and small animals disturb land surfaces with digging,
burrowing, wallowing, trampling, and other behaviors whose effects
may permanently inuence landscape evolution (Butler, 1995). As
expected, large mammals often have very large effects. This paper
aims to describe and discuss the large and small effects of the largest
terrestrial mammal, the African elephant. First I examine some of the
ways in which free-roaming elephants shape and re-shape landscapes
in contemporary African settings. A systematic quantication of the
effects has not yet been reported, but general impacts of the actions
are here summarized. I describe digging, trampling, rock polishing,
and other effects such as habits of destructive feeding on vegetation.
Next I describe fossil sites some containing proboscidean bones
and attempt to explain how ancient sediments preserve traces of
proboscidean behavior very much like the behavior of living
2. Animal landscape-sculptors and ecosystem-engineers
The ways in which terrestrial animals affect surface sediments are
many and varied. Portions of the landscape may be re-shaped and
effectively sculpted by animal behavior. Likewise, some animal
behaviors may change vegetational communities by eliminating,
damaging, or suppressing specic plant taxa.
Beavers (Castor canadensis) provide an example of both processes.
They frequently stack and weave sections of tree branches and sticks
and stabilize them with mud to build dams that create lakes and
ponds, often diverting streamow and altering habitats (see Naiman
et al., 1988 and Butler, 1995: 148183). An extreme example is in
Wood Buffalo National Park, Canada, where beavers have built an 850-
m-long dam in remote, at wetlands within coniferous forest, a
construction feat that probably stretched over many generations
(; accessed 3
September 2010).
An example of smaller-scale animal landscape-sculpting is seen
with modern wolves (Canis lupus), which dig out or enlarge natural
rockshelters and earth hollows, especially in sand, to make dens for
newborn pups (Mech, 1970: 118123). Sometimes these sites are re-
used but also are often abandoned, and other animals then sometimes
adopt them for dens. Wolves are not the only diggers. Warthogs
(Phacocoerus aethiopicus) in Africa modify existing ground openings
such as erosion gullies or antbear burrows, or excavate when needed,
creating shelter from predators at night or protection from heavy rain
(Cumming, 1975). Fossorial rodents tunnel through sediments, some
of which may contain archeological materials, either in open-air or
cave/rockshelter settings, often destroying stratication or creating
features such as stone lines that deceptively appear to be original
strata (Bateman et al., 2003). Sometimes inlled remnants of animal
burrows (called krotovinas) are preserved in ancient sediments (see,
for example, Tappen et al., 2002).
Some animal effects on land surfaces are larger scale. Well-used
animal migration routes following hilltops may become sunken
trailways, such as the bison trails now adopted for highways and
rail routes in the eastern United States. A still noticeable bison trail
system is the Buffalo Trace in Indiana, also called the Vincennes Trace
and Clarksville Trace, where US Route 150 follows it. Another long-
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distance bison trail has become the Natchez Trace Parkway from
Mississippi to Tennessee. Similar animal-made routes are still being
created and used today in southern Africa, where some game trails
have been used by nomadic elephants for centuries (G. Haynes, 1991,
2001, 2005). Fig. 1 shows trails used by elephants (and other species)
in Hwange National Park, Zimbabwe. These trails range from 30 to
50 cm wide and 515 cm in depth, depending on substrate and
intensity of use. Elephant-use creates nearly at-bottomed trails with
fairly regular lateral margins, whereas hoofed animals may create
trails with less even bottoms and margins. Parallel trails may be
created and used in some places, especially when elephants move in
greater numbers. The trails created and maintained by elephants are
compacted and scuffed surcial sediments without covering vegeta-
tion, which has been worn off. Oftentimes fresh and trampled dung
may form a carpeting on sections of the trails (as in Fig. 1 on the left),
especially near water sources where elephant trafc is concentrated.
Laws et al. (1975) recognized that large African animals, such as
elephants, interact with the environment in powerful ways. Currently
researchers are carefully measuring how these interactions have
major organizing effects upon ecosystem processes as well as
structure(McNaughton et al., 1988: 799). Below I describe African
elephant effects on land surfaces and ecosystems that I have observed
or found in the literature.
2.1. Digging by elephants
Elephants are especially able to alter land surfaces on large and
small scales. Elephants in Africa dig wells to reach subsurface water
(Douglas-Hamilton and Douglas-Hamilton, 1975: 165; G. Haynes,
1988, 1991) and excavate into mineral deposits to ingest the
sediments (Buss, 1990: 164170) or to spread dust over their skins
to protect against biting insects and harsh sunlight, leaving behind
large holes and depressions in the ground (Fig. 2; also see gures in
Buss, 1990:166).
Elephants visiting water holes in southern Africa may walk away
from the pond bottoms with mud sticking to their legs and bodies, and
also may ingest some mud, thus deepening and enlarging the water
basins (Weir, 1969). Flint and Bond (1968) estimated that African
elephants in Rhodesia (now Zimbabwe) removed 0.3 to 1.0 m
of mud
every time they wallowed in mud. This process has been documented
for other animals as well in other parts of the world (Butler, 1995).
Elephants in Kenya have been recorded entering a cave system in Mt
Elgon to scrape and feed on minerals from the cave walls in total
darkness, thus reshaping the cave interior (Redmond, 1982). Buss
(1990) found large elephant-excavated holes and pits in roadbanks
and slopes within the Ngorongoro Crater. Analysis of the mineral
samples suggested that the elephants were digging in the sediments
specically for manganese and cobalt, essential micronutrients, and
regularly walked considerable distances to reach these sediments.
2.2. Elephants can smooth and polish rock surfaces
In the Sengwa Wildlife Research Institute, Zimbabwe, elephants
polish rock faces when they press against stone surfaces to drink
water from at least one spring issuing from the base of a bedrock ridge
(Fig. 3). Elephants (and other animals) scratch themselves in African
game reserves by rubbing their bodies against rocks and tree stumps
(Fig. 4), wearing down the stone or wood surfaces and frequently
producing glossy polish.
2.3. Elephants affect vegetational communities
Elephants also affect landscapes in other ways that do not directly
involve sediments, such as stripping bark from live trees for
Fig. 1. Trails used frequently by elephants in Zimbabwe. On the left, a trail in loam showing compression up to 10 cm deep in the center; on the right, a shallower trail in loose
Kalahari sand where the deepening results as much from pushing-up of sand to each side as from compression. Photographed in Hwange National Park; right photograph taken in
late 1990s, left photograph taken 1983.
100 G. Haynes / Geomorphology 157-158 (2012) 99107
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nourishment, or pushing trees over to reach upper branches, often
killing the trees. In Africa, elephants are sometimes considered
problems because they cause the loss of plant species. Any other
animals that may feed on those plants must either migrate or decrease
in numbers as they lose forage sources. Elephants preferentially feed
on certain plants in different seasons of the year, seeking moisture and
nutrients from the tissues. In the dry season, grass which is a favorite
forage in the wet growing seasons dries out and provides poor
forage, so elephants must browse more often, seeking more woody
vegetation, including leaves, twigs, and bark. Heavy feeding on woody
plants may irreversibly damage whole stands of trees and bushes.
According to Sikes (1971: 248), an adult male elephant in Africa may
eat up to 150 kg of vegetable matter every day, but also damages even
more than this daily while feeding (it is a somewhat wasteful
feeder) and moving through vegetation. Elephants are capable of
opening up previously dense vegetation while feeding and trampling.
Higher densities of elephants can completely remove large patches of
woody vegetation, and open up ground for different sorts of
vegetational communities, including patchy mosaics (Lindsay, 1990)
or grasslands. According to Petrides and Swank (1964: 841; cited in
Buss 1990:162), elephants may maintain a relatively early succes-
sional stage of plant community developmentby removing trees,
often enabling other animal species to live in regions where they
would not have been present otherwise. Once wooded habitats that
have been cleared by elephants may experience an increase in animals
that are mixed feeders and grazers. Such changes are major ecological
impacts. Bell (1985) found that woodland dominated by the common
genera Acacia,Commiphora, and Adansonia is particularly affected by
the impacts of elephants, becoming more open and losing consider-
able numbers of trees because of elephant feeding. Such changes in
biota also may affect geomorphological processes, such as fostering
accelerated erosion or slowing the formation of soil horizons.
2.4. Accumulations of elephant dung
Where elephants congregate, such as at preferred feeding patches
or around water sources, dung may be so thick as to carpet the ground
surface over large expanses (Fig. 5). An average size African elephant
may pass up to 100 boluses of 12 kg each, generally in 2030
defecations, every 24 h (Sikes, 1971: 107). Passage time of undigested
food probably varies between about 20 to over 50 h, based on studies
of a captive Asian elephant (Benedict, 1936).
Trampled dung concentrated in aggregation sites, such as margins
of waterholes, would alter local sediment pH near surfaces and clearly
add substantial amounts of organic matter to the mineral sediments.
Measurements of the pH effects of different quantities and densities of
elephant dung are not available, but overall the dung probably does
not dramatically change landscape acidity or alkalinity except on very
localized scales. The process of such heavy deposition of dung is
usually seasonal. At the end of the season of deposition the organic
matter may be overgrown with fresh vegetation and eventually
incorporated into a litter zone until the next episode of dung
deposition and trampling, expectably in a dry season when the
ground cover is once again depleted and minimal. Elephants do not
digest more than about half of what they eat. Because of the 12 day
passage time in the gut the organic matter may be derived from forage
ingested some distance away from the defecation sites; thus,
undigested seeds or nuts may be distributed fairly widely and expand
the range when they successfully germinate.
Fig. 2. Elephant-dug pit in mineral sediments, Hwange National Park, Zimbabwe.
Photographed in late 1990s.
Fig. 3. Smoothed sandstone rock face where elephants (judging from the tracks in the
streambed) and possibly other large mammals have pressed or rubbed against it while
drinking from the spring. Photographed in Chirisa National Park, Zimbabwe, 1982.
Fig. 4. Smoothed and polished stump of Combretum imberbe (leadwood), rubbed by
elephants in Hwange National Park, Zimbabwe. This is a very dense and strong
hardwood, and this degree of polish indicates frequent heavy rubbing by elephants,
suggesting what can also happen with rock surfaces. Photographed in the early 1990s.
101G. Haynes / Geomorphology 157-158 (2012) 99107
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Some undigested materials in dung may be preserved in the
sediments as future macrofossils or as objects that could be mistaken
for human-made materials such as cordage manufactured from plant
bers (Fig. 6).
2.5. Elephants trample and disturb surface materials
A mature male African elephant may weigh over 5000 kg; a
mature female may weigh 3000 kg (Sikes, 1971: Fig. 39, p. 179). The
foot-loadingof a walking modern elephant has been published (see
Guthrie, 1990:263, referring to other sources) as 510660 g/cm
which is the force exerted per unit area of ground surface. This value is
actually lower than the foot-loading of horse or bison, or even the
much smaller saiga antelope, because the weight is spread over a
much broader foot area than in the hoofed taxa. Table 1 shows foot-
loading values for a variety of different northern taxa. So much force is
exerted that each elephant footstep has the power to compact surcial
sediments to a large degree. Studies of the effects of foot-loading from
ungulate and human trampling are numerous in range management
literature (for example, Ferrero, 1991; Mulholland and Fullen, 1991;
Saravi et al., 2005), and show that large-mammal trampling reduces
soil macropore space (impeding water inltration and increasing
surface runoff) and also lowers root biomass and whole plant biomass
for many species, seriously decreasing the fertility index of land
surfaces where trampling is intense. Some plant species may not
survive elephant foot trafc; others may be depressed in growth and
reproduction. Hence, vegetational communities may be greatly
inuenced, especially near water, along habitual trails, or in preferred
feeding patches.
Surcial sediments and plants are not the only features that are
impacted by elephant trampling. Elephant trampling also has a clear
effect on future deposits of fossil bones. Surcial sediment grains may
be abraded against bone surfaces when trodden under the great mass
and large foot size of elephants, creating marks that may be mistaken
for traces of butchering by hominins (Fig. 7). Bones may be marked by
the abrasion, and they may be broken and scattered by kicking or
dragging of elephant feet. These actions surely also affected animal
bones in the distant past, subtracting some bone elements and
modifying others within fossil assemblages.
Elephants often pick up objects to investigate before putting them
down far out of the original place. Elephants are sometimes inclined to
re-arrange the bones of dead elephants encountered at death sites
(see Douglas-Hamilton and Douglas-Hamilton, 1975 for photographs
of live elephants holding and moving bones). The weight of a
trampling proboscidean can fracture animal bones, and on gravel
surfaces the trampling can break stone pebbles and cobbles (Lopinot
and Ray, 2007), creating false artifactssuch as akes and ake-
scarred coresthat have the characteristics of human-made materials.
Some elephant effects, such as digging, trampling, and dust-
bathing, may accelerate erosion of the land-surface or disturb
archeological deposits and invert depositional sequences, confusing
Fig. 5. Dung boluses, trampled dung, and elephant bones on a land surface at a water point, Hwange National Park, Zimbabwe. Photographed around 1983.
Fig. 6. Long woody plant bers deposited in elephant dung, Hwange National Park,
Zimbabwe. Photographed in mid-1980s.
102 G. Haynes / Geomorphology 157-158 (2012) 99107
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future stratigraphic interpretations by mixing materials of very
different ages.
Although many of the inuences that elephants have on
landscapes are localized and relatively small-scale, often they do
have lasting effects on landform shapes and characteristics, recogniz-
able thousands of years after creation. Table 2 summarizes some of the
measurable or estimated effects that African elephants may have on
modern landscapes.
3. Fossil evidence of proboscidean ecosystem engineering and
3.1. Proboscidean effects on extinction
Owen-Smith (1987) proposed that landscape engineering by
prehistoric proboscideans may have fostered the great Pleistocene
diversity seen in non-analog animal communities of the time.
Proboscideans were keystonespecies that partly shaped ecosystems
(a concept taken for granted by many paleoecologists, such as
Robinson et al., 2003), mainly by inuencing the survival or extent
of some vegetational communities, as modern elephants are known to
do. In the past, more sympatric taxa of proboscideans lived in the
world, such as the two or three species of Mammuthus, one species of
Mammut, and several other species of gomphotheres in the Americas.
The cumulative effects of these large mammals kept many habitats in
mosaic-state, with open glades and patches around wooded and
grassy patches, and thus fostered faunal and oral diversity. Human
foragers invaded the proboscidean ranges during the global dispersal
of Homo sapiens after 100 ka, and they may have hunted mammoths
and mastodons to extinction, setting off an ecological cascade of
vegetational changes in North America that led to further extinctions
of many other large herbivores.
The debate continues over the ultimate cause(s) of the end-
Pleistocene extinctions. Owen-Smith's hypothesis about mammoth/
mastodon engineering of local environments is testable, if for example
enough radiocarbon dates from the many taxa that died out actually
do indicate that proboscideans died rst. Abstracts and oral reports
from one as yet inadequately published dating project (Graham et al.,
1997, 2002; Graham, 1998), however, seem to indicate that pro-
boscideans died out last, after all of the other well-dated extinct taxa
had disappeared, in a two-step process which is the opposite of what
Owen-Smith proposed.
3.2. Digging by mammoths
Blackwater Locality Number 1 in New Mexico the original Clovis
archeological site, after which Clovis stone spear points were named
contains spring conduits and wells attributed to human digging,
including some of Holocene age (Green, 1962) and at least one that is
late Pleistocene in age (C. V. Haynes et al., 1999). Well-digging by
humans at the time of Clovis archeological culture is a signicant
event, since a Clovis-age drought (C.V. Haynes, 1991) has been
postulated for much of North America just prior to the end of the
Pleistocene, and may have gured in the process of megafaunal
extinctions. The well at the site is a fairly narrow shaft or circular pit
(C. V. Haynes et al., 1999:455) sunk through underlying sediments
and inlled with sediments from above. It was discovered in 1964 and
re-exposed in 1993. It is interpreted as an unsuccessful hole dug by
Clovis people to reach ground water about 13,500 cal BP (C. V. Haynes
et al., 1999). Fragments of mammoth and bison bones were found
nearby in the archeological excavations. Bear, tapir, badger, and
beaver were ruled out as excavators of the regular cylindrical pit, and
no clear evidence clinches the case that humans dug the pit such as
the presence of artifacts or preserved shovel/scoop marks. It may be
possible that a mammoth dug it using its trunk to grasp bundles of
sediments and throw them aside, as happens in elephant country in
Africa, for example. The 1.5 m depth is within the range seen in
African elephant excavations for water in Zimbabwe (G. Haynes,
3.3. Mammoth tracks
Elephant-foot-size depressions (Fig. 8) on a buried occupation
surface at the Murray Springs archeological site in Arizona, inter-
preted as mammoth tracks (C.V. Haynes, 1973; C.V. Haynes and
Huckell, 2007), were created around 13,000 cal BP. The paleosurface is
thought to be a spring eld where water oozed fromslopesinto a
small stream (C.V. Haynes, 2007: 40). Mammoth bones were found
below and atop the surface, some associated with stone artifacts.
Besides making the tracks in saturated surcial sediments, mam-
moths had also scraped the ground in places within a dry or sluggish
stream bed, possibly in search of water, as elephants do today in Africa
Table 1
A sample of foot-loading values by northern mammalian taxa, (from Guthrie 1990:
262263, citing other sources).
Taxon Foot-loading value (grams per cubic centimeter)
Bison bison bison 10001300
Equus caballus horse 625830
Saiga tatarica saiga antelope 600800
Loxodonta/Elephas elephant 510660
Alces alces moose 420560
Ovibos moschatus musk-ox 325400
Rangifer tarandus caribou 80140
Canis lupus wolf 89114
Homo sapiens R. Dale Guthrie
Fig. 7. Trample marks on a wildebeest (Connochaetes taurinus) tibia, made by elephants.
Photographed in mid-1980s.
103G. Haynes / Geomorphology 157-158 (2012) 99107
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(G. Haynes, 1991). The distinctive type of paleo-surface microrelief at
the site provides a unique window into conditions of the past,
implying drought that may have affected the entire region and
beyond (C.V. Haynes, 1991).
Another fossil site with possible traces made by mammoth feet is
Hot Springs, South Dakota. Nearly 60 mammoths and several
individuals of other vertebrate and invertebrate taxa were trapped
and died in a 20 m deep, steep, slick-sided karst sinkhole that had a
warm spring at the bottom (Agenbroad, 1994; Thompson and
Agenbroad, 2005) over the course of about 350700 years, at 26,000
BP. The sinkhole lled with laminated layers of clay, silt, and sand
around articulated and disarticulated (and unpermineralized) animal
bones. In at least one locus, two sediment proles contain distur-
bances of the horizontally bedded layers, interpreted as mammoth
tracks (Fig. 9).
3.4. Beast solonetzsites
Some Siberian sites rich in mammoth bones appear to have been
frequented by animals in search of mineral sediments. These sites also
may contain abundant evidence of the human presence in the form of
stone tools, used to kill or butcher mammoths at the localities.
Derevianko et al. (2000:53) hypothesized that one suchsite, Shestakovo
in western Siberia, was createdbetween 25,600 to 18,040 BP by humans
seeking mammoths that had been attracted to a local geochemical
landscape, i.e., the solonetz soilwhich contains relatively high levels of
potassium and magnesium. This sort of interpretation has acquired
some traction in the literature. It is becoming better known that there
are so-called beast solonetzlocalities in eastern Europe and parts of
northern Asia mineral deposits visited by mammoths for generations,
where deathsfrequently occurred,and slumping of sediments (possibly
caused by mammoth excavations) sometimes buried bones and
preserved them for the long-term. Leshchinskiy (2001, 2006, 2009)
proposed that Upper Paleolithic humans in northern Eurasia balanced
the search for suitable toolstone against the search for large mammals.
In his view, the largest archeological sites occur where sources of
toolstone are located near mammoth migration routes in landscapes
rich in calcium, magnesium, and sodium. Soffer (1993:40) also
suggested that some central European (Moravian) mammoth-bone
archeological sites, which are often huge and lled with thousands of
bones, were located where they are because of the local mineral-rich
sediments that attracted mammoths. Abraczinskas (1994) tested this
possibility in North America with a spatial analysis of Michigan
mastodont-bone sites and saline water sources, but the mastodon
bone sites did not strongly correlate with the locations of the saline
Table 2
Measurements or rough estimates of elephants' potential geomorphic effects.
Action Quantity or effect Reference
Trampling (foot-loading)
compression force
510660 g per square centimeter Guthrie
Trail creation/use Depth of incision=515 cm or more, depending upon substrate; Width of trail =3050 cm, depending upon substrate
(Note: trails may be created more by compression underfoot than by sediment removal/erosion)
G. Haynes,
unpub. eld
Sediment removed from water holes
after wallowing, carried away on
the body of each individual
0.31.0 Flint and
Dung deposited around water hole
edges at drought refuges
Up to 2+ kg per square meter in dry season, depending upon intensity of use G. Haynes,
1991, and
unpub. eld
Excavation of unconsolidated
sediments (such as well-digging in
sand) by an individual elephant in
one session, using feet and trunk
1 G. Haynes,
unpub. eld
1-season removal of consolidated
mineral sediment by multiple
feeding elephants seeking
13+ cubic meters, depending upon numbers and intensity of feeding G. Haynes,
unpub. eld
Fig. 8. An excavated paleosurface at the Murray Springs archeological site in Arizona,
dated to about 13,000 cal BP (photograph provided by C. V. Haynes, Jr.). These are
interpreted as mammoth footprints around the muddy edge of a pond. A skeleton of an
adult female mammoth (M. columbi) is shown being excavated in the upper part of the
photograph. Photographed in 1966.
104 G. Haynes / Geomorphology 157-158 (2012) 99107
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3.5. Mammoth rub sites
Possible Pleistocene rub sites have been found in North America,
such as at Sonoma Coast State Park in northern California (Parkman,
2002, 2009; Parkman et al., n.d.). These sites are specically
interpreted as places where now-extinct (Rancholabrean) large
mammals scratched themselves against blueschist boulders and
seastacks (Fig. 10), smoothing and polishing some parts of the rock
surfaces (Fig. 11). The polished areas are often located much higher off
the ground than can be reached by modern terrestrial mammals, such
as cattle or horses. The smoothing affected high points on the rock
surfaces rather than entire surfaces, indicating it was a mechanical
process of contact abrasion rather than chemical process such as
weathering or dissolution. The sites are possibly situated on a
migration route that linked coastal prairies with the interior plains
of California. Flaked stone artifacts have been found in excavations at
the Sonoma Coast rub loci, but no direct dates are yet available on
either the cultural materials or the rubbed rock surfaces. Humans
quarrying toolstone from the sites may have removed much of the
once polished outer surfaces of rock (Parkman, 2009). Parkman
(2009) and Peterson (2003) name other possible rub sites in New
Mexico, Nevada, Minnesota, and Wisconsin.
3.6. Mammoth bone accumulations as sediment traps
Loci where elephants died en masse or serially over time from
noncultural causes are fairly uncommon, but enough have been
recorded (for example, see G. Haynes, 1991) to lead me to think that
noncultural sites containing multiple proboscidean skeletons were
also expectably created in the ancient past. Multiple skeletons
accumulate most often near water sources today in Africa, like the
elephant die-off sites in Zimbabwe (G. Haynes, 1991), and waterside
localities probably were also the settings for multiple mammoth/-
mastodon deaths.
Conceivably, large numbers of proboscidean bones would act as
sediment traps and divert streamow and drainage, reshaping land
surfaces and site topography. One possible example is the Colby Clovis
site in Wyoming, dating to about 13,000 cal BP. More than 450 bones
from seven Mammuthus columbi and a few other taxa were found
along with aked stone tools, including spear points probably used to
kill and/or butcher the mammoths. Human stacking of carcass parts or
skeletons along with some redeposition by water created two
concentrated piles of bones and some scattering of others (Frison
and Todd, 1986). Maps of bones (such as Fig. 2.8 in Frison and Todd,
1986, p. 44) and rose diagrams of the excavated bones (Frison and
Todd, 1986: 53) suggest that water owing in the steep sided and
narrow paleo-stream dispersed some elements from the bone piles
that people had made. Experiments done in a modern stream using
recent elephant bones indicate that moving water disperses and also
aggregates bones. Once bones are moved by water, according to Frison
and Todd (1986:64), the potential for subsequent movement is
decreased.Bones may then become dams. When that happens,
scouring and downcutting by water around the clusters or larger
elements would cause bank slumping that might bury bones
episodically (Frison and Todd, 1986: 80).
4. Conclusion
Proboscidean effects on ancient landscapes and archeological
deposits perhaps are underestimated in some settings. A single
trampling elephant can move large amounts of surcial materials just
by the sweep of its feet. Elephant excavations for minerals or water
directly shape parts of landscapes and also inuence erosional effects.
Elephant-abilities to sculpt land surfaces, alter vegetational commu-
nities, and produce mimics of artifacts make proboscideans an
Fig. 9. Sediment disturbances thought to be created by mammoth feet at the Hot
Springs Mammoth Site, South Dakota. Photographed in late 1990s.
Fig. 10. Sea stacksat the Mammoth Rub locus in Sonoma Coast State Park, California. Photographed in 2003.
105G. Haynes / Geomorphology 157-158 (2012) 99107
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unusually important animal to consider in paleogeomorphological
I thank all the generous and hospitable people who encouraged or
supported my work in Africa, especially current and former members
of staff of Zimbabwe's Parks and Wildlife Management Authority.
Their names are too numerous to list anymore. I thank the scientists
and colleagues upon whom I have depended for insights and data,
especially the late Paul Martin, an inspiration in every way, and C.
Vance Haynes, also a true guiding light (and who provided
photographs of the Murray Springs site), along with Larry Agenbroad,
Jeff Saunders, Breck Parkman, George Frison, the late Nikolai
Vereshchagin, Alexei Tikhonov, Vadim Garutt, Piotr Wojtal, Kate
Scott, the late Antony Sutcliffe, Tony Stuart, Adrian Lister, and dozens
more. I am most beholden to Janis Klimowicz.
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... They are known to be keystone species-species that have a highly significant effect on community structure relative to their biomass or abundance-involved in modification of vegetation structure (e.g. Bakker et al., 2016;Haynes, 2012), dispersal of seeds (e.g. Campos-Arceiz and Blake, 2011; Janzen and Martin, 1982), and recycling and spreading of nutrients (e.g. ...
... Elephants also often push over trees to feed on the higher parts of the foliage. These destructive behaviours can transform woody vegetation into grasslands (Haynes, 2012). Like other megaherbivores, elephants ate fruits and contributed to the dispersion of the seeds (Campos-Arceiz and Blake, 2011). ...
... Both woolly mammoths and the straight-tusked elephants, like their extant relatives, were probably able to dig wells to reach subsurface water or excavate the soil to ingest the sediments for their mineral deposits (Haynes, 2012). They certainly also strongly modified the landscape by trampling and by the formation of trails that were used repeatedly as migration routes. ...
Dietary traits of individuals and populations of both Neanderthals and animals, are essential for the reconstruction of biotic interactions among species. These kinds of dynamic relationships with other living species in a shared environment can be seen as a major influence in evolution and ecology, and the timing and type of interaction could have driven many aspects of Neanderthal behaviour. This chapter is aimed at combining the different approaches to studying interactions between organisms or populations: dietary food webs of plants-animals (stable isotopes and tooth wear), animals-humans (stable isotopes) and more complex interactions between humans and other animals, mainly carnivores (based on taphonomy and zooarchaeology).
... This points to a high concentration of alkaline earth and alkali elements in the ground and pore waters of sediments that form the geochemical landscape of VG. These elements-primarily Ca, Mg, and Na-and some trace elements create the basis for favorable geochemical landscapes, which are very attractive to large herbivores of past and present times, in particular, representatives of the Proboscidea (Cooper, 1831;Leshchinskiy, , 2012Leshchinskiy, , 2015Walker et al., 2001;Holdø et al., 2002;Mwangi et al., 2004;Haynes, 2012). At the end of the Pleistocene, mineral starvation caused by chronic geochemical stress led to the concentration of large mammals around the beast solonetz. ...
... Modern elephants often move the bones and tusks of deceased relatives, especially in places of their concentration (e.g., near watering sites and on mineral licks). They also transform the microrelief by digging large pits and even caves using tusks and limbs during their search for mineral-containing sediments (Haynes, 2012). This behavior probably also was typical for mammoths, which could greatly alter the primary structure of thanatocoenosis and stratigraphy at faunal localities. ...
This paper describes the results of research at Volchia Griva, the largest site in Asia containing mammoth fauna in situ. It is situated in the south of the West Siberian Plain in the Baraba forest-steppe zone, and occupies an area of several hectares. Analysis of sediments and taphonomy of the site allows us to suggest that thousands of megafaunal remains were buried here in mud pits and erosional depressions. The favorable geochemical landscape of Volchia Griva attracted animals during periods of mineral starvation. This is reflected in the high mortality in two intervals, ca. 20–18 ¹⁴ C ka BP and ca. 17–11 ¹⁴ C ka BP. The results of palynological analysis of samples from the upper part of the Volchia Griva section made it possible to reconstruct the history of landscape changes of the Baraba Lowland during the MIS 2. Forb-mesophytic meadows were common at the beginning of this period, with taiga type forests. At ca. 20 ¹⁴ C ka BP, an abrupt and significant aridization of the climate occurred, which led to the degradation of forests. The mammoth steppe was widely developed, dominated by forb-grass association and with areas of alkali meadows and soils. Such conditions existed probably until the mid-Holocene.
... There were 330 species of herbs and 120 shrubs recorded on granite bedrock, with corresponding figures of 316 and 114, respectively, for basalts. In terms of frequency, the most common species were grasses Panicum maximum (recorded in 54, i.e., 90% of all plots), Brachiaria deflexa (52), Tragus berteronianus (49), and Urochloa mosambicensis (43), herbs Phyllanthus maderaspatensis (47), Hibiscus micranthus (41), and Acalypha indica (40), and shrubs Flueggea virosa (46), Dichrostachys cinerea (44), and Philenoptera violacea (39). In total, 29 taxa occurred in more than half of the plots sampled (Table 1). ...
... The effect of herbivores, elephants in particular, on vegetation has been thoroughly studied 49 , yet not in the perspective of the two factors addressed in our study or directly exploring the role of seasonal rivers. With high population densities, elephant herds influence the savanna ecosystem not only by consuming large amounts of plant tissue (mainly leaves) but also by damaging or uprooting grown trees 18,50 , hence changing woody savanna into more open grass dominated states 51,52 . While some plant species cope with these disturbances well, other important tree canopy species regenerate poorly 53 . ...
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To identify factors that drive plant species richness in South-African savanna and explore their relative importance, we sampled plant communities across habitats difering in water availability, disturbance, and bedrock, using the Kruger National Park as a model system. We made plant inventories in 60 plots of 50 × 50 m, located in three distinct habitats: (i) at perennial rivers, (ii) at seasonal rivers with water available only during the rainy season, and (iii) on crests, at least~ 5 km away from any water source. We predicted that large herbivores would utilise seasonal rivers’ habitats less intensely than those along perennial rivers where water is available throughout the year, including dry periods. Plots on granite harboured more herbaceous and shrub species than plots on basalt. The dry crests were poorer in herb species than both seasonal and perennial rivers. Seasonal rivers harboured the highest numbers of shrub species, in accordance with the prediction of the highest species richness at relatively low levels of disturbance and low stress from the lack of water. The crests, exposed to relatively low pressure from grazing but stressed by the lack of water, are important from the conservation perspective because they harbour typical, sometimes rare savanna species, and so are seasonal rivers whose shrub richness is stimulated and maintained by the combination of moderate disturbance imposed by herbivores and position in the middle of the water availability gradient. To capture the complexity of determinants of species richness in KNP, we complemented the analysis of the above local factors by exploring large-scale factors related to climate, vegetation productivity, the character of dominant vegetation, and landscape features. The strongest factor was temperature; areas with the highest temperatures reveal lower species richness. Our results also suggest that Colophospermum mopane, a dominant woody species in the north of KNP is not the ultimate cause of the lower plant diversity in this part of the park.
... By contrast, rates of bioturbation (vertical soil mixing) are inversely related to body size [39], and animal-induced bioturbation was recently shown to be significant across a range of biomes [40]. While large mammals, such as suids (e.g., wild boar), proboscideans (e.g., elephants), and equids (e.g., donkeys), can make considerable contributions to bioturbation in certain ecosystems [41][42][43][44], associated fossorial mammals and soil macrofauna are generally more effective bioturbation agents [28]. ...
... Soil mixing and processing by associated fauna Large animals can expose a larger fraction of the organic matter in soils to organo-mineral interaction through vertical soil mixing, either through their own activity [41][42][43][44] or the activity of associated fossorial mammals or soil fauna [28,82,83] (Figure 1). The highest known soil mixing rates are reported in open or semi-open ecosystems historically associated with large herds of herbivores (savannas and grasslands [40]), but we lack good quantification of soil mixing rates in many systems. ...
There is growing interest in aligning the wildlife conservation and restoration agenda with climate change mitigation goals. However, the presence of large herbivores tends to reduce aboveground biomass in some open-canopy ecosystems, leading to the possibility that large herbivore restoration may negatively influence ecosystem carbon storage. Belowground carbon storage is often ignored in these systems, despite the wide recognition of soils as the largest actively-cycling terrestrial carbon pool. Here, we suggest a shift away from a main focus on vegetation carbon stocks, towards inclusion of whole ecosystem carbon persistence, in future assessments of large herbivore effects on long-term carbon storage. Failure to do so may lead to counterproductive biodiversity and climate impacts of land management actions.
... As an example, while the elephants' repeated use of migration trails or paths leading to water sources might have facilitated humans in the practice of particular hunting strategies (Haynes, 2012;Agam and Barkai, 2018), it might have as well triggered intensive trampling especially in those places where accumulations of elephant carcasses usually occur. As a consequence of trampling and kicking (by elephants and other megaherbivores), stratigraphy may be reworked, bones and artifacts may be dispersed and reoriented, edge-damages may occur on stone tools and marks and fractures may be variably produced on bones to the extent of mim-icking cut-marks or intentional breaking (Fiorillo, 1984;Andrews and Cook, 1985;Gifford-Gonzalez et al., 1985;Behrensmeyer et al., 1986;Haynes, 1988Haynes, , 2012Olsen and Shipman, 1988;Nielsen, 1991;Domínguez-Rodrigo et al., 2009;Gaudzinski-Windheuser et al., 2010;McPherron et al., 2014;Courtenay et al., 2019aCourtenay et al., , 2020Pizarro-Monzo and Domínguez-Rodrigo, 2020). ...
... As an example, while the elephants' repeated use of migration trails or paths leading to water sources might have facilitated humans in the practice of particular hunting strategies (Haynes, 2012;Agam and Barkai, 2018), it might have as well triggered intensive trampling especially in those places where accumulations of elephant carcasses usually occur. As a consequence of trampling and kicking (by elephants and other megaherbivores), stratigraphy may be reworked, bones and artifacts may be dispersed and reoriented, edge-damages may occur on stone tools and marks and fractures may be variably produced on bones to the extent of mim-icking cut-marks or intentional breaking (Fiorillo, 1984;Andrews and Cook, 1985;Gifford-Gonzalez et al., 1985;Behrensmeyer et al., 1986;Haynes, 1988Haynes, , 2012Olsen and Shipman, 1988;Nielsen, 1991;Domínguez-Rodrigo et al., 2009;Gaudzinski-Windheuser et al., 2010;McPherron et al., 2014;Courtenay et al., 2019aCourtenay et al., , 2020Pizarro-Monzo and Domínguez-Rodrigo, 2020). Since direct evidence of elephant trampling, such as ichnofossils, are rarely preserved (Palombo et al., 2018;Serangeli et al., 2020), inferences are substantially drawn from the indirect evidence. ...
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Human-proboscidean interactions are key nodes of complex ecological, cultural and socio-econom- ic systems. In the last decades, evidence has been provided in support of an early human exploitation of proboscidean carcasses, offering further insights into past human behaviors, diet and subsistence strategies. Nevertheless, the mode of acquisition of the carcasses, the degree of exploitation, its timing relative to carnivore scavenging and to the decom- position of the carcass, its ecological and socio-eco- nomical role are hitherto not fully understood and a matter of debate. By summarizing the empirical evidence for human-elephant interactions in Early and Middle Pleistocene open-air sites of western Eurasia, this contribution elaborates on the need for a more rigorous, spatially explicit inferential procedure in modeling past human behaviors. A renewed analytical approach, namely spatial ta- phonomy, is introduced. In its general term, spatial taphonomy refers to the multiscale investigation of the spatial properties of taphonomic processes. Building upon a long lasting tradition of tapho- nomic studies, it seeks for a more effective theoret- ical and methodological framework that accounts for the spatio-temporal dimension inherent to any complex system. By bridging into a spatio-tempo- ral framework the traditional archaeological, geo- archaeological and taphonomic approaches, spatial taphonomy enhances our understanding of the processes forming archaeological and palaeonto- logical assemblages, allowing a finer comprehen- sion of past human behaviors.
... Behaviour such as geophagy can also affect an animal's movement. Geophagy is the consumption of essential minerals from mineral licks (Bowell et al., 1996;Holdø et al., 2002;Ruggiero and Fayz, 1994;Weir, 1969), and is thought to also have been practiced by mammoths (Haynes, 2006(Haynes, , 2012Leshchinskiy, 2012Leshchinskiy, , 2017Soffer, 1993;Zenin et al., 2006). These factors, however, rarely prompt elephants to travel tens of kilometres. ...
The ecology and behaviour of woolly and Columbian mammoths and mastodons have been extensively studied. Despite this, their patterns of mobility, and particularly the question of whether or not they migrated habitually, remains unclear. This paper summarises the current state of knowledge regarding mobility in these species, reviewing comparative datasets from extant elephant populations as well as isotopic data measured directly on the ancient animals themselves. Seasonal migration is not common in modern elephants and varies between years. Nonetheless, non-migratory elephants can still have considerable home ranges, whose size is affected mainly by habitat, seasonal availability of water and food, and biological sex. Strontium isotope analyses of woolly mammoths, Columbian mammoths, and mastodons demonstrate plasticity in their migratory behaviour as well, probably in response to spatio-temporal variations in ecological conditions. However, biological sex is difficult to establish for most proboscidean fossils and its influence on the results of Sr analyses can therefore not be assessed. Advances in intra-tooth sampling and analytical methods for strontium isotope analysis have enabled research on intra-annual movement, revealing nomadic behaviour in all three species. Sulfur isotopes have been analysed from woolly mammoth remains numerous times, but its methodology is not yet developed well enough to inform on past proboscidean mobility in as much detail as strontium studies. The inter- and intra-individual variation in migratory behaviour in mammoths and mastodons implies that their role in the subsistence strategies of Palaeolithic people may have fluctuated as well. Further assessment of hominin-proboscidean predator-prey interactions will require a more detailed understanding of proboscidean habitual mobility in specific contexts and places. Strontium isotope studies based on multi-year enamel sequences from multiple individuals have the potential to provide this insight.
... African savannah elephants (Loxodonta africana), hereafter referred to as "elephants", are mega-herbivores well known for their role in ecosystem engineering [8][9][10]. Elephants perform important ecological roles, such as affecting the quality of foliage available through their foraging activities [11]. ...
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African savannah elephants (Loxodonta africana) are well-known as ecosystem engineers with the ability to modify vegetation structure. The present study aimed to examine how male elephant foraging behaviour is affected across (a) season (wet versus dry); (b) time of day (before or after noon); (c) presence or absence of other elephants; and (d) reproductive state (musth versus no musth). Six radio-collared adult elephant bulls were observed twice per week from June 2007–June 2008 in Kruger National Park (KNP), South Africa. Using generalized linear mixed effect modeling, results indicate that elephant bulls graze more during the wet season and browse more during the dry season. To potentially offset the costs associated with thermoregulation during the heat of the day, KNP elephants spent more time foraging during the morning, and more time resting during the afternoon. Male elephants also foraged significantly less when they were associated with females compared to when they were alone or with other males. This is likely due to male–female associations formed mainly for reproductive purposes, thus impeding on male foraging behaviours. In contrast, the condition of musth, defined by the presence of related physical signs, had no significant effect on foraging behaviour.
ZoogeomorphologyZoogeomorphology is a relatively new discipline and is set for changes this century in BotswanaBotswana that have not been seen for millennia. The co-evolution of landscapes and wildlife amidst a changing climate means that the precise role of each is difficult to determine. As southern Africa becomes hotter and drier due to anthropogenic climate change this century, past adaptation strategies such as wildlife movements along (Balinsky’s 1962) ‘drought corridor’ will no longer be possible due to land use/land cover change and agriculture and human-related expansion. As the Sixth Great Extinction unfolds, it offers a unique opportunity to study just how significant different biota are in determining geomorphology, albeit as our climate changes. FaunaFauna that has remained intact since the Miocene will largely disappear from African shores and the palaeo-duneDune fields of the Kalahari may well become reactivated. The real significance of micro-organisms and invertebrates will then be realised as the bioturbators remain to shape the landscapes around them without the distraction posed by humans and the unique mega-fauna that surrounded them.
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The crisis facing Africa’s elephant populations is a notorious example of ongoing wildlife declines caused by illegal harvesting. Targeted conservation interventions require detailed knowledge about changes in population sizes and the effect of illegal activities. However, accurately quantifying poaching intensity is a difficult task: commonly calculated from ranger-based carcass-encounter data, the proportion of illegally killed elephants (PIKE) is a function of poaching and background mortality. Hence, at constant poaching intensity, PIKE decreases with increasing natural mortality and also with hunting, management interventions, and other anthropogenically induced deaths. Natural mortality is often more difficult to quantify with accuracy than mortality due to illegal killing, as elephants that die naturally are more likely to be missed than those taken by poachers. In recent analyses, constant background mortality rates were assumed. Yet, for example climate-driven fluctuations in natural mortality, if not quantified and accounted for, may lead to biased estimates of poaching intensity. Varying background mortality rates can be accounted for in the analysis of PIKE, but this requires near-complete counts of natural and management-related deaths and hunting records. Carefully developed population models, which simulate population dynamics and demographic changes while accounting for variation in environmental conditions and management strategies, are alternatives. However, successful calibration of such models requires integrating comprehensive demographic data. We systematically review the scientific and ‘grey’ literature on African elephant demography with the objective of facilitating poaching and population analysis possibilities through an inventory of information relevant to demography. Our screening of 10900 publications resulted in the review of relevant information provided by 431 studies from 420 study sites throughout Africa. From these, we extracted demographic data collected between 1900 and 2017, and collated them in the newly created African Elephant Demographic Database (AEDD; 10.6084/m9.figshare.19387085). We found 37 natural mortality estimates from five different study sites. Other mortality data, demographic rates, and age- and sex-structured population data were substantially more abundant, both temporally and spatially. This new collection of demographic rates, age- and sex-structured population data, and cause-partitioned mortality estimates identifies spatial and temporal data gaps and provides prior information needed for African elephant population models. Closing these data gaps and subsequent analyses of realistic population models may aid elephant conservation via improved policies, legislation, and protection.
Conference Paper
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African elephants are vital to their ecosystems, but their populations are threatened by a rise in human-elephant conflict and poaching. Monitoring population dynamics is essential in conservation efforts; however, tracking elephants is a difficult task, usually relying on the invasive and sometimes dangerous placement of GPS collars. Although there have been many recent successes in the use of computer vision techniques for automated identification of other species, identification of elephants is extremely difficult and typically requires expertise as well as familiarity with elephants in the population. We have built and deployed a web-based platform and database for human-in-the-loop re-identification of elephants combining manual attribute labeling and state-of-the-art computer vision algorithms, known as ElephantBook. Our system is currently in use at the Mara Elephant Project, helping monitor the protected and at-risk population of elephants in the Greater Maasai Mara ecosystem. ElephantBook makes elephant re-identification usable by non-experts and scalable for use by multiple conservation NGOs.
The Eastern Gravettian technocomplex is a widespread entity that dominated Central and Eastern Europe between some 28,000 and 10,000 B.P. (Kozlowski 1986, 1990; Otte 1982). This poorly understood entity contains within it a number of archaeological cultures that show affinities to each other and are classified into the Willendorf, Pavlov, Kostenki, and Avdeevo cultures and grouped into various larger-sized cultural entities. The reasons for the similarities between them are poorly understood and generally involve postulations about some sort of a west-to-east population movement (Grigor’ev 1968; Kozlowski 1986, 1990; Tarasov 1979). Here I argue that a new understanding of these archaeological entities can be gained by embedding them into the natural world in which the hunter-gatherers who made the inventories functioned, specifically by considering the one species, mammoth, that played a significant role in their adaptations.