Sample And Methods

We obtained information on body mass (kg) and RMR (kcal/day) for 41 primate species, including 17 species of strepsirrhine and 24 species of haplorhine from published sources, from which we calculated a single unweighted average for each species (Table 1). All RMR values are expressed as kilocalories per day (kcal/day) and were converted from other units when necessary.

Data on brain mass (g) and body mass (kg) for 15 strepsirrhine species and 21 haplorhines were obtained from Bauchot and Stephan (1969) and Stephan et al. (1981). For each species, we calculated a single unweighted average for both brain mass and body mass (Table 2). Humans were excluded from the

1 We follow Kurland and Pearson (1986) in defining hypometabolism as having a RMR more than 20% below that predicted for body size by the Kleiber scaling relationship. This conservative definition is used in order to avoid the misclassification of a species as hypometabolic as a result of measurement error, the measurement of an animal during sleep, or due to lack of standardized procedures.

2 Kurland and Pearson (1986) used the traditional Prosimii-Anthropoidea taxonomic split but, since they did not include Tarsius in their analysis, there is no difference between their use of prosimian and our use of strepsirrhine.

Table 1. Sample information for primate metabolic and ecological data

Species Metabolic data® Ecological data4

RMR Body Deviation5 DQ^ Habitat' Activity

(kcal/day) mass (kg) cycled

Suborder Strepsirrhini

Arctocebus calabarensis

15.2

0.206

-28.99

327.5

A

N

Cheirogaleus medius

22.7

0.300

-20.00

-

A

N

Eulemur fulvus

42.0

2.397

-68.85

129.0

A

D

Euoticus elegantulus

25.1

0.260

-1.52

230.0

A

N

Galago moholi

13.9

0.155

-19.62

-

A

N

Galago senegalensis

18.1

0.215

-18.11

278.0

A

N

Galagoides demidoff

6.3

0.058

-23.85

305.0

A

N

Lemur catta

45.1

2.678

-69.22

166.0

A

D

Lepilemur ruficaudatus

27.6

0.682

-47.46

149.0

A

N

Loris tardigradus

14.8

0.284

-45.65

327.5

A

N

Microcebus murinus

4.9

0.054

-37.51

-

A

N

Nycticebus coucang

32.4

1.380

-63.65

-

A

N

Otolemur crassicaudatus

47.6

0.950

-29.33

195.0

A

N

Otolemur garnettii

47.8

1.028

-33.13

275.0

A

N

Perodicticus potto

41.3

1.000

-41.00

190.0

A

N

Propithecus verreauxi

86.8

3.080

-46.67

200.0

A

D

Varecia variegata

69.9

3.512

-61.08

-

A

D

Suborder Haplorhini

Alouatta palliata

231.9

4.670

+4.28

136.0

A

D

Aotus trivirgatus

52.4

1.020

-26.25

177.5

A

N

Callithrix geoffroyi

27.0

0.225

+18.07

235.0

A

D

Callithrix jacchus

22.8

0.356

-29.23

235.0

A

D

Cebuella pygmaea

1G.1

0.105

-21.78

249.5

A

D

Cercopithecus mitis

407.7

8.500

+17.00

201.5

T

D

Cercocebus torquatus

196.2

4.000

-0.90

234.0

A

D

Colobus guereza

357.9

10.450

-12.03

126.0

A

D

Erythrocebus patas

186.9

3.000

+17.13

-

T

D

Homo sapiens

1400.0

53.500

+1.10

-

T

D

Hylobates lar

123.4

1.900

+8.93

181.0

A

D

Leontopithecus rosalia

51.1

0.718

-6.41

-

A

D

Macaca fascicularis

400.9

7.100

+31.67

200.0

T

D

Macaca fuscata

485.4

9.580

+27.34

223.0

T

D

Macaca mulatta

231.9

5.380

-6.22

159.0

T

D

Pan troglodytes

581.9

18.300

-6.05

178.0

T

D

Papio anubis

342.9

9.500

-9.47

207.0

T

D

Papio cynocephalus

668.9

14.300

+29.95

184.0

T

D

Papio papio

297.3

6.230

+7.70

-

T

D

Papio ursinus

589.3

16.620

+2.27

189.5

T

D

Pongo pygmaeus

569.1

16.200

+0.68

172.5

A

D

Saguinus geoffroyi

5G.5

0.500

+21.43

263.0

A

D

Saimiri sciureus

68.8

0.850

+11.03

323.0

A

D

Tarsius syrichta

8.9

0.113

-34.80

350.0

A

N

"McNab and Wright (1987); Leonard and Robertson (1994); Thompson et al. (1994); Kappeler (1996).

^Richard (1985); Sailer et al. (1985); Nowak (1991); Napier and Napier (1994); Rowe (1996). 'Metabolic deviation from predicted by Kleiber equation. ^Dietary quality.

A = primarily arboreal; T = primarily terrestrial. -D = diurnal; N = nocturnal.

Table 2. Sample information for primate brain data

Species

Brain mass (g)a

Body mass (kg)a

Suborder Strepsirrhini

Arctocebus calabarensis

7.2

0.323

Cheirogaleus medius

3.1

0.177

Eulemur fulvus

25.2

2.397

Euoticus elegantulus

7.2

0.274

Galago senegalensis

4.8

0.186

Galagoides demidoff

3.4

0.081

Lemur catta

25.6

2.678

Lepilemur ruficaudatus

7.6

0.682

Loris tardigradus

6.6

0.322

Microcebus murinus

1.8

0.054

Nycticebus coucang

12.5

0.800

Otolemur crassicaudatus

10.3

0.850

Perodicticus potto

14.0

1.150

Propithecus verreauxi

26.7

3.480

Varecia variegata

34.2

3.512

Suborder Haplorhini

Alouatta palliata

51.0

6.400

Aotus trivirgatus

16.0

0.850

Callithrix geoffroyi

7.6

0.280

Callithrix jacchus

7.6

0.280

Cebuella pygmaea

4.5

0.140

Cercopithecus mitis

76.0

6.500

Cercocebus torquatus

104.0

7.900

Colobus guereza

73.0

7.000

Erythrocebus patas

118.0

8.000

Hylobates lar

102.0

6.000

Macaca fascicularis

74.0

5.500

Macaca fuscata

84.0

5.900

Macaca mulatta

110.0

8.000

Pan troglodytes

420.0

46.000

Papio anubis

205.0

26.000

Papio cynocephalus

195.0

19.000

Papio papio

190.0

18.000

Papio ursinus

190.0

18.000

Pongo pygmaeus

370.0

55.000

Saguinus geoffroyi

10.0

0.380

Saimiri sciureus

22.0

0.680

»Bauchot and Stephan (1969); Stephan et al. (1981).

»Bauchot and Stephan (1969); Stephan et al. (1981).

analysis because they are outliers for brain size in relation to body size and consequently substantially alter regressions. Because of differences between the body masses of animals used for brain studies and those for metabolic studies, when comparing metabolic rates to brain size, we calculated an adjusted RMR for each species to account for this difference.

Information on dietary quality (DQ) was obtained for 12 strepsirrhine and 20 haplorhine species (Table 1) from Richard (1985), Rowe (1996) and Sailer et al. (1985). Diet quality was assessed using an index, developed by Sailer et al. (1985), which considers the relative energy and nutrient density of dietary items. The DQ index is a weighted average of the proportions of foliage, reproductive plant material, and animal material. The DQ is calculated as:

Here s = percent of diet derived from structural plant parts (e.g., leaves, stems, and bark), r = percent of diet derived from reproductive plant parts (e.g., fruits, flowers, nectar, and resin), and a = percent of diet derived from animal parts (including both vertebrates and invertebrates). The DQ ranges from a minimum of 100 (100% foliage) to a maximum of 350 (100% animal material). Humans were excluded from the dietary analysis because the range of possible diets is larger than any nonhuman primate species, and consequently an all-inclusive DQ for the human species is not possible.

To assess functional consequences of substrate and habitat use, we classified species as arboreal or terrestrial based on primary habitat (Table 1); this determination was derived from relevant literature (Nowak, 1991; Rowe, 1996). While this dichotomy is overly simplified, it is used simply to get a general picture of habitat use. Additionally, we obtained information on activity cycle (i.e., nocturnal or diurnal) from published sources for all 17 species of strepsirrhine and all 24 haplorhine species (Rowe, 1996; Table 1).

To examine the evolutionary context of RMR in primates, we compiled metabolic data for closely related mammalian orders. We obtained information on RMR (kcal/day) and body mass (kg) for bats (order Chiroptera) and tree shrews (order Scandentia) from published sources, from which we calculated a single unweighted average for each species (Table 3). No metabolic data were available for colugos (order Dermoptera). All RMR values are expressed as kilocalories per day (kcal/day) and were converted from other units when necessary.

We compiled data on body mass (kg) estimates for 16 species of subfossil Malagasy lemurs from Godfrey et al. (1997; Table 4). Body mass reconstructions, based on regressions of humeral and femoral midshaft circumferences

Table 3.

J. Josh Snodgrass et al. Sample information for RMR and body mass for selected mammalian species

Species RMR (kcal/day)a Body mass (kg)"

Order Chiroptera

Anoura caudifer

4.07

0.012

Artibeus lituratus

9.82

0.070

Carollia perspicilla

3.64

0.015

Chalinolobus gouldii

2.92

0.018

Chrotopterus auritus

11.80

0.096

Cynopterus brachyotis

5.45

0.037

Desmodus rotundus

3.06

0.029

Diaemus youngi

3.99

0.037

Diphylla ecaudata

3.96

0.028

Dobsonia minor

12.71

0.087

Eonicterus spelaea

5.61

0.052

Glossophaga longirostris

3.07

0.014

Glossophaga soricina

2.50

0.010

Hipposideros galeritus

1.08

0.009

Histiotus velatus

1.16

0.011

Leptonycteris curasoae

3.95

0.024

Leptonycteris sanborni

5.10

0.022

Macroderma gigas

10.94

0.107

Macroglossus minimus

2.39

0.016

Megaloglossus woermanni

2.52

0.012

Miniopterus schreibersii

3.01

0.011

Molossus molossus

4.61

0.056

Noctilio albiventris

2.75

0.027

Noctilio leporinus

5.44

0.061

Nyctimene albiventer

4.64

0.028

Nyctophilus geoffroyi

1.32

0.008

Nyctophilus major

2.36

0.014

Paranyctimene raptor

3.36

0.021

Phyllostomus discolor

4.06

0.034

Phyllostomus elongatus

4.55

0.036

Phyllostomus hastatus

8.18

0.084

Pteropus poliocephalus

36.74

0.598

Pteropus scapulatus

28.12

0.362

Rhinonycteris aurantius

1.88

0.008

Rhinophylla fisherae

1.88

0.010

Rousettus aegyptiacus

14.22

0.146

Sturnira lilium

4.56

0.022

Syconycteris australis

3.92

0.018

Tonatia bidens

4.48

0.027

Uroderma bilobatum

3.08

0.016

Order Scandentia

Ptilocercus lowii

5.04

0.058

Tupaia glis

10.84

0.123

»Bradley and Hudson (1974); Whittow and Gould (1976); McNab (1988); Arends et al. (1995); Geiser et al. (1996); Hosken (1997); Hosken and Withers (1997, 1999); Bartels et al. (1998); Baudinette et al. (2000).

Table 4. Reconstructed body masses (kg) and cranial capacities (cc) for selected subfossil Malagasy lemur species

Species Body mass (kg)® Cranial capacity (cc)b

Table 4. Reconstructed body masses (kg) and cranial capacities (cc) for selected subfossil Malagasy lemur species

Species Body mass (kg)® Cranial capacity (cc)b

Family Archaeolemuridae

Archaeolemur edwardsi

22.0

104 c,i

Archaeolemur majori

17.0

Hadropithecus stenognathus

28.0

Family Daubentoniidae

Daubentonia robusta

10.0

Family Lemuridae

Pachylemur insignis

10.0

Pachylemur jullyi

12.5

46c

Family Megaladapidae

Megaladapis edwardsi

75.0

Megaladapis grandidieri

65.0

Megaladapis madagascariensis

40.0

118'

Family Palaeopropithecidae

Archaeoindris fontoynontii

200.0

Babakotia radofilai

15.0

49d

Mesopropithecus dolichobrachion

12.0

Mesopropithecus globiceps

10.0

Mesopropithecus pithecoides

11.0

Palaeopropithecus ingens

45.0

Palaeopropithecus maximus

55.0

99c

»Godfrey et al. (1997). ^Ravosa (unpublished data). ^British Museum (Natural History). dDuke University Primate Center.

»Godfrey et al. (1997). ^Ravosa (unpublished data). ^British Museum (Natural History). dDuke University Primate Center.

indicate that the subfossil lemurs were all larger than living strepsirrhine primates. Some species had body masses slightly greater than the largest living strepsirrhines (Indri indri and Propithecus diadema) (Smith and Jungers, 1997); however, all known species appear to have had body masses of at least 10 kg (Godfrey et al., 1997). Numerous species were considerably larger, including Archaeoindris fontoynontii, which is estimated to have reached an adult mass of 200 kg. We additionally present data on cranial capacity (cc) for five species of subfossil Malagasy lemur, which were collected by M. Ravosa (unpublished data) (Table 4).

Allometric relationships were determined using ordinary least squares regressions (OLS)3 of log10-transformed data. Additionally, allometric relationships were calculated using reduced major axis (RMA); however, RMA values are not reported because they were not significantly different from parameters calculated using OLS. Differences in regression parameters were assessed using Student's t-tests. All analyses were performed using SPSS (Version 8.0), except RMA equations, which were calculated using BIOMstat (Version 3.30a).

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