Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ELECTRONIC SUPPLEMENTARY MATERIAL
16
14
Vertebrates (10)
Arthropods (54)
Plants (7)
Microbes and Fungi (4)
Soil nutrients (9)
Number of studies
12
10
8
6
4
2
0
1990
1995
2000
2005
2010
ESM Fig. 1. Summary of the 84 case studies, from 65 papers, found using ISI Web of
Science that examined the effects of carrion on five different ecological categories between
1990 and 2010. Some studies examined multiple categories (e.g. soil nutrients and plant
responses).
1
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
30
Multiple types
25
Number o studies
Large carcasses
20
Small carcasses
15
Pig
10
5
0
Farmland
Forest
Grassland
Urban
Arid
ESM Fig. 2. Summary of carrion arthropod studies across five major habitat types using
different animal carrion models. ‘Farmland’ included modified production landscapes;
‘Forest’ included forest and woodland environments; ‘Grassland’ included prairie and
savannah environments; ‘Urban’ included built environments; ‘Arid’ included desert and
semi-arid environments. ‘Small carcasses’ = fish, rodents, rabbits or birds; ‘Large carcasses’
= ungulates or large vertebrates. ‘Multiple types’ = two or more different animal carcasses.
Some studies contributed to more than one category (e.g. compared arthropods at rodent and
rabbit carcasses in farmland and nearby forest). All studies presented empirical data on the
response of arthropods to carrion between 1990 and 2010. List of studies are shown in Table
S4.
2
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
30
Three or more taxa
25
Number o studies
Diptera and Coleoptera
20
Coleoptera only
15
Diptera only
10
5
0
Habitats
Seasons
Carcass
type
None
ESM Fig. 3. Summary of arthropod studies that included a spatial or temporal contrast, and
focused on a particular arthropod group. ‘Habitats’ included spatial environmental
comparisons; ‘Seasons’ included within or between year comparisons; ‘Carcass types’
included comparisons of carrion from different animal species; ‘None’ included studies with
no spatial or temporal contrast. Some studies contributed to more than one category (e.g.
compared coleopteran communities in different habitats at different times of the year). All
studies presented empirical data on the response of arthropods to carrion between 1990 and
2010. List of studies are shown in Table S4.
3
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ESM Table 1. Summary of the duration and depth to which carcasses affect soil nitrogen
(NH4+, NO3-, or total N).
Carcass type
Duration
Soil
Depth
Biome
Reference
Bison, Cattle, Whitetailed deer
36 months
10cm
Tall-grass prairie, USA
(Towne 2000)
Red kangaroo
2 months
5cm
Southern arid zone,
Australia
(Wilson and Read
2003)
Cicadas
3 months
10cm
Deciduous forest, USA
(Yang 2004)
Bison
12 months
10cm
Pig
100 days
5cm
White-tailed deer
Moose
15 months for
soil
15 months for
soil
10cm
Temperate lowland forest,
Poland
Temperate woodland,
Canada
Temperate hardwood
forest, USA
10cm
Boreal forest, USA
Mule deer
39 months
46cm
Shrub-steppe, USA
Dog
39 months
38cm
Shrub-steppe, USA
White-tailed jackrabbit
27 months
18cm
Shrub-steppe, USA
Uinta ground squirrel
27 months
21cm
Shrub-steppe, USA
Least chipmunk
15 months
15cm
Shrub-steppe, USA
Magpie
15 months
15cm
Shrub-steppe, USA
Sage sparrow
15 months
15cm
Shrub-steppe, USA
Terrestrial garter snake
15 months
15cm
Shrub-steppe, USA
Northern leopard frog
27 months
15cm
Shrub-steppe, USA
(Melis et al. 2007)
(Benninger et al.
2008)
(Bump et al. 2009b)
(Bump et al. 2009a)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
(Parmenter and
MacMahon 2009)
4
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ESM Table 2. Summary of studies on microbial or fungal response to carrion.
Carcass type
Duration
Microbial response
Spatial
contrast
Biome
Reference
Cicadas
1 month
elevated bacterial
and fungal diversity
Carcass vs
control
Deciduous
forest, USA
(Yang 2004)
Rats
28 days
elevated microbial
biomass
None
Tropical
savanna,
Australia
(Carter et al.
2008)
Moose
40 months
elevated bacterial
and fungal diversity
Carcass vs
control
boreal forest,
USA
(Bump et al.
2009a)
Pig
71 days
altered bacterial
community
None
Tropical urban
grassland,
USA
(Howard et al.
2010)
5
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ESM Table 3. Summary of plant responses to carrion.
Carcass
type
Muskox
whitetailed deer
Duration
Plant
response
>5 yrs
biomass
2yrs
biomass, foliar
nitrogen
Moose
2 yrs
foliar nitrogen
Bison
Bison,
cattle,
whitetailed deer
5 yrs
foliar nitrogen
3 yrs
Cicadas
10 weeks
biomass,
species
composition
seed mass,
foliar nitrogen
Cicadas
2 yrs
herbivory
Contrast
Distance
from
carcass
Carcass vs
control
Carcass vs
control
Carcass vs
control
Distance
from
carcass
Carcass vs
control
Carcass vs
control
Biome
Reference
Arctic
temperate
hardwood forest,
USA
(Danell et al. 2002)
boreal forest, USA
temperate lowland
forest, Poland
(Bump et al. 2009a)
tall-grass prairie,
USA
Deciduous forest,
USA
Deciduous forest,
USA
(Bump et al. 2009b)
(Melis et al. 2007)
(Towne 2000)
(Yang 2004)
(Oliver et al. 2004)
6
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ESM Table 4. Summary of ecological studies on arthropod communities associated with
carrion for the period 1990-2010.
Carrion type
Arthropod
groups
Study
duration
Spatial or temporal
contrasts
Biome
Reference
Pig
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera
Diptera
70 days
none
semidesert,
Mexico
2 years
Seasons and forest
type
Among seasons
Diptera,
Coleoptera,
Hymenoptera
Coleoptera
90 days
12 months
between surface and
burrow, among
seasons
Among seasons
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera,
Hymenoptera,
Isoptera
Diptera,
Coleoptera
Diptera,
Coleoptera
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera
Coleoptera
2 years
Among seasons
3 months
none
Temperate
forest, Poland
Subtropical
rural farmland,
Argentina
semi-arid
shrub-steppe,
USA
Pine woodland,
Turkey
Urban
Guangzhou,
China
Sugarcane
crop, Brazil
(ValdesPerezgasga et
al. 2010)
(Matuszewski
et al. 2010)
(Battan
Horenstein et
al. 2010)
(Parmenter and
MacMahon
2009)
(Ozdemir and
Sert 2009)
(Shi et al.
2009)
3 years
River catchments
Coastal Pacific
north, Canada
(Hocking et al.
2009)
30 days
Open savanna vs
shaded cloud forest
Montane
Venezuala
(Velasquez
2008)
57 days
Forest types
25 weeks
shaded vs open,
among seasons
Carcass type,
between seasons
Temperate
forest, Poland
Prairie, Canada
(Matuszewski
et al. 2008)
(Sharanowski
et al. 2008)
(Moretti et al.
2008)
Diptera,
Coleoptera,
Hymenoptera,
Araneae,
Hemiptera
Pig
Pig
Rat
Pig
Rabbit
Pig
Salmon
Rat
Pig
Pig
Mouse and
Rat
Pig
Fish
Pig
1 year
11 months
1 year
Between seasons
2 months
Between habitats
2 weeks
none
Subtrpical
secondary
forest, Brazil
subtropical
urban, China
Temperate
forest,
grassland,
urban, Poland
Tropical Palm
oil plantation,
Malaysia
(Gomes et al.
2009)
(Wang et al.
2008)
(Ulrich et al.
2008)
(Chin et al.
2007)
7
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
Carrion type
Arthropod
groups
Study
duration
Spatial or temporal
contrasts
Biome
Reference
Fish
Coleoptera
2 months
Between habitats
(Ulrich et al.
2007)
Chicken
Coleoptera
8 days
Between road types
Pig
Diptera,
Coleoptera
2 months
none
Possums
Diptera
15 days
oviposition
exclusion/opportunity
Pig
Diptera
14 months
Between habitats
Bear, Deer
Alligator, Pig
Diptera,
Coleoptera,
Hymenoptera,
Araneae,
Hemiptera,
Acari,
Lepidoptera,
Blattodea
Diptera,
Coleoptera
Diptera,
Coleoptera,
Hymenoptera,
Isopoda, Acari
Diptera,
Coleoptera
Coleoptera
73 days
Between seasons
Temperate
forest,
grassland,
urban, Poland
Deciduous
temperate
forest, USA
Tropical
mountain,
Colombia
Grazed
temperate
grassland,
Australia
Urban London,
England
Mixed
flatwood
forest, USA
21 days
Between seasons
60 days
none
Farmland,
USA
urban Vienna,
Austra
(Tabor et al.
2005)
(Grassberger
and Frank
2004)
21 days
Between seasons
26 days
Carcass vs. control
Diptera,
Coleoptera,
Hymemoptera,
Araneae,
Acari,
Collembola
Coleoptera
1 year
Between seasons
Farmland,
USA
Boreal forest,
Norway
Mediterranean
farmland,
Spain
(Tabor et al.
2004)
(Melis et al.
2004)
(Arnaldos et al.
2004)
40 days
Between meadow
and deciduous orest
(Kocarek 2003)
Diptera,
Coleoptera,
Hymenoptera,
Isoptera
Diptera,
Coleoptera
12 days
Between sites
Temperate
grassland and
forest, Czech
Republic
Subtropical
woodland and
farmland, USA
2 years
among seasons
temperate
sclerophyll
forest,
Australia
(Archer 2003)
Pig
Pig
Pig
Roe deer
Chicken
Rat
Pig
Pig
(Dunn and
Danoff-Burg
2007)
(Martinez et al.
2007)
(Lang et al.
2006)
(Hwang and
Turner 2005)
(Watson and
Carlton 2005)
(Shahid et al.
2003)
8
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
Carrion type
Arthropod
groups
Study
duration
Spatial or temporal
contrasts
Biome
Reference
Pig
Diptera,
Coleoptera
2 years
among seasons
(Archer and
Elgar 2003)
Pig
Diptera
6 days
none
Rabbit
Diptera,
Coleoptera,
Hymenoptera,
Araneae,
Solpugidae
Diptera,
Coleoptera,
Hymenoptera,
Araneae,
Hemiptera,
Acari,
Lepidoptera,
Blattodea
Diptera
30 months
none
temperate
sclerophyll
forest,
Australia
temperate
sclerophyll
forest,
Australia
Tropical urban,
India
92 days
Between carcass type
Subtropical
flatwood
forest, USA
(Watson and
Carlton 2003)
3 days
Between habitats
15 days
Between habitats
(Woodcock et
al. 2002)
(Gibbs and
Stanton 2001)
Chicken
Diptera,
Coleoptera,
Acari,
Opiliones
Diptera
Woodland,
Scotland
Temperate
mixed forest,
USA
1 year
Between seasons
(Arnaldos et al.
2001)
Cow
Diptera
2 years
Between habitats
Rabbit
Diptera,
Coleoptera
Diptera,
Coleoptera
2 months
Between habitats
16 months
Diptera,
Coleoptera,
Hymenoptera
Diptera,
Coleoptera
50 days
Above vs below
ground, between
habitats
Between habitats
Mediterranean
farmland,
Spain
Mediterranean
woodland,
Spain
Coastal dunes,
France
Coastal and
sub-boreal,
Canada
Montane and
alpine, USA
Bear, Deer
Alligator, Pig
Slugs
Chicken
Pig
Rabbit
Pig
1 month
Coleoptera
Pig
Pig
Diptera,
Coleoptera
Diptera,
Coleoptera
1 month
271 days
(Archer and
Elgar 2003)
(Bharti and
Singh 2003)
between habitats,
between carcass type
Upland forest,
Hawaii
(MartinezSanchez et al.
2000)
(Bourel et al.
1999)
(VanLaerhoven
and Anderson
1999)
(De Jong and
Chadwick
1999)
(Avila and Goff
1998)
Between habitats,
between seasons
Between habitats,
between seasons
open vs sunlit
southeast Spain
(Pinero 1997)
Upland forest,
Hawaii
Rural
farmland,
Canada
(Richards and
Goff 1997)
(Anderson and
VanLaerhoven
1996)
9
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
Carrion type
Arthropod
groups
Study
duration
Spatial or temporal
contrasts
Biome
Reference
Rabbit
Diptera,
Coleoptera
6 months
Between seasons
(Tantawi et al.
1996)
Rat, Rabbit,
Goat
Diptera
10 months
Between habitats,
between seasons
urban
Alexandria,
Egypt
Savanna,
Texas, USA
Rat
Diptera,
Coleoptera
between habitats
(Wells and
Greenberg
1994)
(Kentner and
Streit 1990)
10
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
ESM Table 5. Summary of studies on vertebrate responses to carrion.
Carcass
type
Measurement
European
bison
diversity of
scavengers
carrion
removal rate,
diversity of
scavengers
Elk
carrion
removal,
Brown hare
Scavenger
type
Temporal
duration
Spatial
contrast
Biome
Reference
avian
5 days
between
carcasses
Patagonia,
Argentina
(Travaini et al.
1998)
avian,
mammal,
240 days
between
carcasses
avian,
mammal,
3.5 years
Wolf
predation
Lowland
temperate
forest, Poland
Meadow and
forest,
yellowstone
NP, USA
12 days
Between
habitats
Temperate
upland pine
and
bottomland
hardwood
forest, USA
5 days
Scavenger
exclusion
House mice
and brown
rats
carrion
removal rate,
diversity of
scavengers
Red
kangaroo
carrion
removal rate
avian,
mammal
avian,
mammal,
reptile
Ungulates
diversity of
scavengers
avian,
mammal,
Various
carrion
removal rate
avian and
mammal
1000 days
Between
habitats
Carcass
type,
scavenger
exclusion
carrion
removal rate
avian
15 days
Between
habitats
diversity of
scavengers
avian and
mammal
16 days
Between
habitats
Barbary
sheep
Horses and
European
bison
1500 days
(Selva et al.
2003)
(Wilmers et al.
2003)
(DeVault et al.
2004)
Southern arid
zone, Australia
Lowland
temperate
forest, Poland
(Read and
Wilson 2004)
semi-arid
shrub-steppe,
USA
Mediterranean
woodland,
Spain
Lowland
temperate
forest, Poland
(Parmenter and
MacMahon
2009)
(Selva et al.
2005)
(Blazquez et al.
2009)
(CortesAvizanda et al.
2009)
11
Barton, Cunningham, Lindenmayer, Manning. (2012). The role of carrion in maintaining
biodiversity and ecological processes in terrestrial ecosystems. Oecologia.
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