Jeschke

Electronic Supplementary material to Jeschke:When carnivores are “full and lazy”

Kill Rates of Free-Ranging Carnivorous Birds and Mammals Compared to their Demands

Carnivore species / Season / Kill rate KR / DEMAND / NKR<DEMAND/Ntotal
Birds
Accipiter gentilis
(northern goshawk) / Year-round
Winter / ♂♂: >202 g/d, ♀♀: >270 g/d (1)
♂♂: 133 g/d, ♀♀: 189 g/d (2) / ♂♂: 108-132 g/d, ♀♀: 144-176 g/d (2) / 0/4
Falco peregrinus
(peregrine falcon) / Breeding / 188 g/d (3) / 128-156 g/d (4) / 0/1
Phalacrocorax carbo
(great cormorant) / Breeding / ♂♂: 787, 969, 1017 g/d (5)
♀♀: 516, 810, 1230 g/d (5) / 747-913 g/d (6)
668-816 g/d (6) / 1/6
Stercorarius parasiticus (Arctic skua) / Breeding / 45 g/d, 47 g/d (7) / 121-147 g/d (8) / 2/2
Mammals
Acinonyx jubatus (cheetah) / ? / ≈4.0 kg/d (9) / 3.1-3.9 kg/d (10) / 0/1
Canis lupus (wolf) / Year-round
Winter
? / 5.6 kg/d (11); 8.2, 10, 14, 17 kg/d (12)
3.1, 3.9, 4.9, 5.7, 5.9, 6.5, 6.5, 7.2, 7.5, 7.6, 7.7 kg /d (13); 5.4 kg/d (14); 6.8 kg/d (15); 6.1, 17.1 kg/d (16); 8.1, 10, 12, 13 kg/d (17)
1.1, 12, 13, 16, 18, 18, 19, 22, 23, 24, 24, 27, 34, 36 kg/d (18) / 2.5-3.1 kg/d (19) / 1/38 (3%)
Canis latrans (coyote) / Winter / 0.50, 0.53, 0.66, 0.94, 1.1, 1.4, 1.9, 3.0 kg/d (20) / 0.66-0.80 kg/d (21) / 2/8 (25%)
Crocuta crocuta
(spotted hyena) / Winter-spring
Winter
Summer-fall
? / 3.8 kg/d (22)
2.0 kg/d (23)
4.0 kg/d (24)
3.8 kg/d (9), 3.8 kg/d (25) / 3.6-4.4 kg/d (26) / 1/5 (20%)
Hyaena brunnea
(brown hyena) / ? / ≈3.2 kg/d (9) / 2.8-3.4 kg/d (27) / 0/1
Lutra lutra (Eurasian otter) / Fall / >1 kg/d (28) / 0.65-0.79 kg/d (29) / 0/1
Lycaon pictus
(African wild dog) / Year-round / 2.7 kg/d (30) / 1.8-2.2 kg/d (31) / 0/1
Lynx canadensis
(Canadian lynx) / Year-round
Winter
? / >0.72 kg/d (32)
0.84, 0.85, 1.0, 1.1, 1.3, 1.3, 1.5,1.6 kg/d (20); 0.39, 0.56, 0.63, 0.96, 1.1 kg/d (33)
1.0, ≈1.3 kg/d (34) / 0.53-0.65 kg/d (35) / 1/16 (6%)
Lynx lynx (Eurasian lynx) / Year-round / Kittens: ≈1.9 kg/d (36)
Subadults: 1.7 kg/d (36)
Adult ♂♂: 3.4 kg/d (36), 3.6 kg/d (37)
Adult ♀♀: ≈1.9 kg/d (37), 3.4 kg/d (36) / 0.72-0.88 kg/d (38)
1.1-1.3 kg/d (38)
1.2-1.4 kg/d (38)
1.5-1.9 kg/d (38) / 0/6
Panthera leo (lion) / Year-round
Winter-spring
?
?
? / Adult ♀♀: 5.0, 5.3, 7.6 kg/d (39)
Adults: 5.9 kg/d (22)
Adults: 4.9, 5.1 kg/d (40)
Adult ♂♂: 7.0, 7.2 kg/d (40)
Adult ♀♀: 4.7, 5.0 kg/d (40) / 4.4-5.4 kg/d (41)
5.5-6.7 kg/d (41)
5.5-6.7 kg/d (41)
6.6-8.0 kg/d (41)
4.4-5.4 kg/d (41) / 2/10 (20%)
Panthera pardus (leopard) / Year-round / Adult ♂♂: 3.5 kg/d (42) / 3.2-3.9 kg/d (43) / 0/1
Puma concolor (cougar) / Year-round
? / Adults: ≈3.9, ≈10.2 kg/d (44)
Adult ♂♂: 8.6 kg/d (45)
Adult ♀♀: 7.1 kg/d (45)
Adults: ≈6.7, ≈8.3 kg/d (46) / 2.8-3.5 kg/d (45)
2.2-2.7 kg/d (45)
3.5-4.2 kg/d (45)
2.8-3.5 kg/d (45) / 0/6

Notes: For a general outline of the calculation of KR and DEMAND, see the main body text. Each DEMAND is given as a range, calculated as the estimated mean value 10%.

(1)Kenward (1977)

(2)Kenward et al. (1981)

(3)Redpath and Thirgood (1997)

(4)Nagy et al. (1999) gave a FMR of 341 kJ/d for a 211 g weighing kestrel (Falco tinnunculus). According to Dunning (1993), adult male peregrines weigh on average 611 g and females weigh 952 g, giving with FMR M0.681 for birds a FMR of 703.4 kJ/d for males and 951.4 kJ/d for females, thus 827 kJ/d for an average adult peregrine. Because of Jensen’s inequality (Ruel and Ayres 1999), one should calculate the FMR for each sex separately and use these values to calculate the FMR averaged over both sexes. Taking just the body mass averaged over both sexes instead leads to a slight systematic overestimation of FMR. The PE is here 0.0129 (Humphreys 1979), giving a DEMAND of 142 g/d.

(5)Grémillet et al. (2004)

(6)Nagy et al. (1999) gave an allometric equation for the FMR of an average marine bird: 14.25 M0.659 kJ/d, giving with M = 3200 g for male and 2600 g for female great cormorants in Greenland (Grémillet et al. 2004) a FMR of 2908.8 kJ/d for males and one of 2536.8 kJ/d for females. Since the caloric value of cormorants’ prey in Greenland is known and markedly lower than the average energy content of animals (4.2 kJ/g compared to 7 kJ/g), I use this value to calculate DEMAND. The PE is here 0.0129 (Humphreys 1979), giving a DEMAND of 830 g/d and 742 g/d for males and females, respectively.

(7)Caldow and Furness (1991, 2001) observed Arctic skuas (Stercorarius parasiticus) during the breeding seasons of 1986 and 1987. There, the skuas forage only during daylight hours (Caldow and Furness 1991). Since Caldow and Furness (2001) observed the skuas at a latitude of 60° N and a longitude of 2° W between mid-June and mid-August, there were about 20 daylight hours per day. Skuas forage when they are off-territory, and off-territory times in 1986-1987 for skua pairs with one or two chicks, respectively, have been given by Caldow and Furness (1991) in their Table 1:

In case of skua pairs with one chick, one adult was off-territory 26% of the available foraging time (5.2 h/d) in the first 8 days post hatching and 51% (10.2 h/d) of the rest of the pre-fledging stage. This latter period is about 20 days because the total fledging period is about 28 days (RSPB 2004). Over the total fledging period, one adult was hence off-territory on average about 5.2 h/d  8/28 + 10.2 h/d  20/28 = 8.8 h/d. When foraging, one skua stole on average 1.7 fish/h from auks. I calculated this value from the raw data underlying Caldow and Furness (2001) which were kindly provided by Richard Caldow. The average consumption rate was thus 15 fish/d. Caldow (pers. comm.) reported a mean fish length of 11.6 cm (N = 40) what converts to an average fresh mass of 6.6 g. The average consumption rate was hence 99 g-fish/d. According to Phillips (2001), fish represent about 94% of the diet of Arctic skuas in Shetland. This value yields a total average consumption rate of 105 g/d that had to be divided between two adults and one chick. Assuming that the chick got 34% the amount of food an adult got (Drent et al. 1992) gives a consumption of 45 g/d per adult.

In case of skua pairs with two chicks, one adult was off-territory 34% of the available foraging time (6.8 h/d) in the first 8 days post hatching and 59% (11.8 h/d) of the rest of the pre-fledging stage. Over the total fledging period, one adult was thus off-territory about 6.8 h/d  8/28 + 11.8 h/d  20/28 = 10.4 h/d, giving an average consumption rate of 18 fish/d and a total of 126 g/d for both adults and chicks and one of 47 g/d per adult.

(8)Nagy et al. (1999) gave an allometric equation for the FMR of an average marine bird: 14.25 M0.659 kJ/d, giving with M = 405 g for male and 467 g for female Arctic skuas in Shetland (Phillips 2001) a FMR of 745.0 kJ/d for males and one of 818.3 kJ/d for females, thus an average FMR of 782 kJ/d. The PE is here 0.0129 (Humphreys 1979), giving a DEMAND of 134 g/d.

(9)Reviewed by Henschel and Tilson (1988)

(10)Nagy et al. (1999) gave an allometric equation for the FMR of an average species of the taxon Carnivora: 1.67 M0.869 kJ/d, giving with M = 55000 g for males and 45000g for females (Estes 1991) a FMR of 21983.2 kJ/d for males and one of 18465.4 kJ/d for females, thus an average FMR of 20224 kJ/d The PE is here 0.0318 (Humphreys 1979), giving a DEMAND of 3.5 kg/d.

(11)Jędrzejewski et al. (2002)

(12)Calculations as in (18) but based on data from Hayes and Harestad (2000).

(13)Calculations based on data from Dale et al. (1994) for wolves preying on caribou (Rangifer tarandus caribou). M (caribou) was set to 65 kg which is about the average mass of an adult caribou (86 kg according to Smith et al. 2003) multiplied by 0.75, a rough correction for the fact that wolf do not only consume adults.

(14)Carbyn (1983)

(15)Hebblewhite et al. (2003)

(16)Mech et al. (2001)

(17)Calculations based on data from Smith et al. (2004) for wolves preying on elk (Cervus elaphus). M (elk) was set to 164 kg (218 kg [Smith et al. 2003]  0.75).

(18)Messier (1994) reviewed wolf kill rates on moose (Alces alces) in which the latter represented > 75% of wolf ungulate diets. To estimate total wolf kill rate, I multiplied the kill rates given by Messier by 1.2. M (moose) was set to 270 kg (359 kg [Smith et al. 2003]  0.75).

(19)Nagy et al. (1999) gave a FMR of 17,700 kJ/d for a 37.3 kg weighing wolf. The average adult North American wolf mass is according to Smith et al. (2003) 42.8 kg, giving with FMR M0.869 for Carnivora (Nagy et al. 1999) a FMR of 15,955 kJ/d. The PE is here 0.0314 (Humphreys 1979), giving a DEMAND of 2.8 kg/d.

(20)Calculations based on data from O’Donoghue et al. (1998) for coyotes or lynx, respectively, preying on snowshoe hares (Lepus americanus). M (snowshoe hare) was set to 1.3 kg (1.71 kg [Smith et al. 2003]  0.75).

(21)Laundré and Hernández (2003) gave an Edemand of 5144 kJ/d what translates to 0.73 kg/d (see main body text).

(22)Green et al. (1984)

(23)Kruuk (1972)

(24)Henschel and Tilson (1988)

(25)Henschel and Skinner (1990)

(26)Calculations as for cheetah but with M = 54000 or 63150 g for males or females, respectively (Estes 1991), giving a DEMAND of 4.0 kg/d.

(27)Calculations as for cheetah but with M = 42700 or 43900 g for males or females, respectively (Estes 1991), giving a DEMAND of 3.1 kg/d.

(28)Carss et al. (1990)

(29)Calculations as for cheetah but with M = 8000 g (Carss et al. 1990), giving a DEMAND of 0.72 kg/d.

(30)Estes and Goddard (1967)

(31)Nagy et al. (1999) gave a FMR of 15,300 kJ/d for a 25.17 kg weighing wolf. The average mass of the African wild dogs studied by Estes and Goddard (1967) was about 18 kg, giving with FMR M0.869 for Carnivora (Nagy et al. 1999) a FMR of 11,500 kJ/d. The PE is here 0.0314 (Humphreys 1979), giving a DEMAND of 2.0 kg/d.

(32)Calculations based on data from Saunders (1963) for lynx preying on snowshoe hares.

(33)Brand et al. (1976)

(34)Calculations based on data reviewed by O’Donoghue et al. (1998) for lynx preying on snowshoe hares.

(35)According to O’Donoghue et al. (1998), the demand of Canadian lynx equals about 0.45 hares/d, i.e. 0.59 kg/d.

(36)Jobin et al. (2000)

(37)Okarma et al. (1997)

(38)Calculations as for cheetah but with M = 9000, 15000, 15700, or 21200 g for an average kitten, subadult, adult female, or adult male lynx (Okarma et al. 1997), respectively, giving a DEMAND of 0.80, 1.2, 1.3, or 1.7 kg/d, respectively.

(39)Van Orsdol (1986)

(40)Reviewed by Green et al. (1984)

(41)Edemand of free-ranging lions is about 270 kJ/kg/d (reviewed by Green et al. 1984), giving with M = 158, 126, or 189 kg for an average adult, female, or male, respectively (Estes 1991), a DEMANDof 6.1, 4.9, or 7.3 kg/d, respectively.

(42)Bothma and le Riche (1986)

(43)Calculations as for cheetah but with M = 50000 (Estes 1991), giving a DEMAND of 3.5 kg/d.

(44)Reviewed by Ackerman et al. (1986)

(45)Ackerman et al. (1986)

(46)Calculations based on data reviewed by Ackerman et al. (1986) for cougars preying on mule deer (Odocoileus hemionus) and porcupines (Erithizon dorsatum). M (mule deer) was set to 58 kg (Ackerman et al. 1986) and M (porcupine) to 5.3 kg (7.1 kg [Smith et al. 2003]  0.75).

References

Ackerman BB, Lindzey FG, Hemker TP (1986)Predictive energetics model for cougars. In: MillerSD, Everitt DD (eds) Cats of the world: biology, conservation and management. National Wildlife Federation, Washington, DC, pp 333-352

Bothma J du P, le Riche EAN (1986)Prey preference and hunting efficiency of the Kalahari Desert leopard. In: Miller SD, Everitt DD (eds)Cats of the world: biology, conservation and management. National Wildlife Federation, Washington, DC, pp 389-414

Brand CJ, Keith LB, FischerCA (1976)Lynx responses to changing snowshoe hare densities in central Alberta. JWildlMgmt 40:416-428

Caldow RWG, Furness RW (1991) The relationship between kleptoparasitism and plumage polymorphism in the arctic skua Stercorarius parasiticus (L.). Funct Ecol 5:331-339

Caldow RWG, Furness RW (2001)Does Holling’s disc equation explain the functional response of a kleptoparasite?J Anim Ecol 70:650-662

Carbyn LN (1983)Wolf predation on elk in Riding Mountain National Park, Manitoba. JWildl Mgmt 47:963-976

CarssDN, Kruuk H, Conroy JWH (1990)Predation on adult Atlantic salmon, Salmo salar L., by otters, Lutra lutra (L.), within the River Dee system, Aberdeenshire, Scotland. J Fish Biol 37:935-944

Drent RH, Klaassen M, Zwaan B (1992)Predictive growth budgets in terns and gulls. Ardea 80:5-17

Dunning JB (ed) (1993)CRC handbook of avian body masses. CRC Press, Boca Raton, FL

Estes RD (1991) The behavior guide to African mammals. University of California Press, Berkeley, CA

Estes RD, Goddard J (1967) Prey selection and hunting behavior of the African wild dog. JWildl Mgmt 31:52-70

Green B, Anderson J, Whateley T (1984)Water and sodium turnover and estimated food consumption in free-living lions (Panthera leo) and spotted hyaenas (Crocuta crocuta). JMammal 65:593-599

Grémillet D, Kuntz G, Delbart F, Mellet M, Kato A, Robin J-P, Chaillon P-E, Gendner J-P, Lorentsen S-H, Le Maho Y (2004)Linking the foraging performance of a marine predator to local prey adundance. Funct Ecol 18:793-801

Hayes RD, HarestadAS (2000)Wolf functional response and regulation of moose in the Yukon. Can JZool 78:60-66

Hebblewhite M, Paquet PC, Pletscher DH, Lessard RB, Callaghan CJ (2003)Development and application of a ratio estimator to estimate wolf kill rates and variance in a multiple-prey system. Wildl Soc Bull 31:933-946

Henschel JR, Skinner JD (1990)The diet of the spotted hyaenas Crocuta crocuta in KrugerNational Park. Afr JEcol 28:69-82

Henschel JR, Tilson RL (1988)How much does a spotted hyaena eat? Perspectives from the Namib Desert. AfrJEcol 26:247-255

Humphreys WF (1979) Production and respiration in animal populations. J Anim Ecol 48:427-453

Jędrzejewski W, Schmidt K, Theuerkauf J, Jędrzejewska B, Selva N, Zub K, Szymura L (2002) Kill rates and predation by wolves on ungulate populations in Białowieża primeval forest (Poland). Ecology 83:1341-1356

Jobin A, Molinari P, Breitenmoser U (2000)Prey spectrum, prey preference and consumption rates of Eurasian lynx in the Swiss Jura Mountains. Acta Theriol 45:243-252

Kenward RE (1977)Predation on released pheasants (Phasanius colchicus) by goshawks (Accipiter gentilis) in Central Sweden. Viltrevy 10:79-109

Kenward RE, Marcström V, Karlbom M (1981)Goshawk winter ecology in Swedish pheasant habitats. JWildl Mgmt 45:397-408

Kruuk H (1972)The spotted hyena. University of Chicago Press, Chicago, IL

Laundré JW, Hernández L (2003)Total energy budget and prey requirements of free-ranging coyotes in the Great Basin desert of the western United States. JArid Environ 55:675-689

Mech LD, Smith DW, Murphy KM, MacNulty DR (2001)Winter severity and wolf predation on a formerly wolf-free elk herd. JWildl Mgmt 65:998-1003

Messier F (1994)Ungulate population models with predation: a case study with the North American moose. Ecology 75:478-488

Nagy KA, GirardIA, Brown TK (1999)Energetics of free-ranging mammals, reptiles, and birds. Annu RevNutr 19:247-277

O’Donoghue M, Boutin S, Krebs CJ, Zuleta G, Murray DL, Hofer EJ (1998)Functional responses of coyotes and lynx to the snowshoe hare cycle. Ecology 79:1193-1208

Okarma H, Jędrzejewski W, Schmidt K, Kowalczyk R, Jędrzejewska B (1997) Predation of Eurasian lynx on roe deer and red deer in Białowieża Primeval Forest, Poland. Acta Theriol 42:203-224

Phillips RA (2001) Stercorarius parasiticus Arctic skua. BWP Update 3:25-41

Redpath SM, Thirgood SJ (1997)Birds of prey and red grouse. Stationary Office, London

RSPB (2005) Arctic skua.

Saunders JK (1963)Food habits of the lynx in Newfoundland. JWildl Mgmt 27:384-390

Smith DW, Drummer TD, Murphy KM, Guernsey DS (2004)Winter prey selection and estimation of wolf kill rates in Yellowstone National Park, 1995-2000. JWildl Mgmt 68:153-166

Smith FA, LyonsSK, Ernest SKM, Jones KE, Kaufman DM, Dayan T, Marquet PA, Brown JH, Haskell JP (2003)Body mass of late Quaternary mammals. Ecology 84:3403

Van Orsdol KG (1986) Feeding behaviour and food intake of lions in Rwenzori National Park, Uganda. In: Miller SD, EverittDD (eds) Cats of the world: biology, conservation and management. National Wildlife Federation, Washington, DC, pp 377-388