J. Zool., Lond.

(2004) 262, 145159

C 2004 The Zoological Society of London

Printed in the United Kingdom

DOI:10.1017/S0952836903004485

Limb proportions and adaptations in caviomorph rodents

(Rodentia: Caviomorpha)

A. Elissamburu and S. F. Vizcano

Departamento Cientfico Paleontologa de Vertebrados; Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, CONICET, Argentina
(Accepted 1 July 2003)

Abstract
Caviomorph rodents (Rodentia: Caviomorpha) are an anatomically variable group with diverse ecological and
morphological types, including cursorial, digging, fossorial and swimming forms. Their appendicular skeleton is
rather generalized and extreme modifications or reductions in the long bones of limbs are not present. It is of
special interest to evaluate functional adaptive variations of the appendicular skeleton within this group. Although
much work has been devoted to study structure, function, and kinematics of legs in mammals, most studies do not
consider particular functions and behaviour within different mammalian lineages. Morphometric and biomechanical
studies were performed to test the relationship between adaptation and morphology of the bones of the proximal
and middle limb segments in terrestrial caviomorph rodents. Fifty-four specimens belonging to 10 species with
different limb adaptations and representing a wide range of body sizes were measured. Diameters and functional
lengths of long bones were taken and nine functional indices were built. Humerus (HRI) and ulna (URI) robustness,
humerus deltoid (SMI) and epicondyle (EI) development, olecranon proportion (IFA), femur (FRI) and tibia (TRI)
robustness, and gluteal muscle insertion at femur (GI), were calculated. Only TRI and, to a lesser degree, SMI
and EI, were significantly related to body mass. A functional sequence (cursorialgeneralizedoccasional digging
diggers) which seems to reflect an increase in force and muscular development in middle segments of the fore
limb is recognized. The hind limb shows a decrease in the speed efficiency of the femur and an increase in limb
robustness in the transition from cursorial to digging forms. Although overlapping of speed and force functions in
the limbs is evident, functional differentiation for speed in the proximal, and force in the middle segments can be
inferred.
Key words: caviomorph rodents, limb proportions, multivariate analyses, biomechanics, adaptation

INTRODUCTION

Caviomorph rodents (Rodentia: Caviomorpha) are a characteristic group of South American mammals that
diversified during the geographic isolation of the continent
during part of the Cenozoic era. They are first recorded
in the Late Oligocene, and developed a large variety of
adaptive types, greater than that of any equivalent rodent
taxon (Vucetich & Verzi, 1999).
In contrast to other rodent groups, living caviomorphs
vary widely in body size (< 200 g in Ctenomys to
> 50 kg in Hydrochaeris), and show great anatomical and
ecological diversity. They include forms specialized for
cursoriality, similar to small ungulates (e.g. Dolichotis),
digging (e.g. Lagostomus), fossorial (e.g. Ctenomys),
scansorial (e.g. Coendu), swimming (e.g. Myocastor), and
large cursorial and swimming forms (e.g. Hydrochaeris).

E-mail: elissamburu@hotmail.com
vizcaino@museo.fcnym.unlp.edu.ar (if 1st address not accepted)

Despite this variability, their appendicular skeleton is

rather generalized and rodents do not show the extreme
modifications of the long bones with reduction of the
digits, as seen in the Artiodactyla and Perissodactyla.
They do, however, show some adaptations convergent
with other mammals adapted to cursoriality (with
predominance of speed function), such as ungulates,
carnivores, lagomorphs, or digging (with predominance
of force function) as in other rodents, and xenarthrans.
Convergences are mainly expressed in the structure and
disposition of the limb segments.
Caviomorph rodents studied in this paper (Table 1)
can be placed into six groups based on behavioural
and morphological features: (1) cursorial (terrestrial
quadrupeds that possess vertically oriented limbs that
move in a parasagittal plane; Stein & Casinos, 1997);
(2) generalized (non-cursorial without digging activity);
(3) occasional diggers (non-cursorial with some digging
activity); (4) diggers (non-cursorial with frequent digging
activity); (5) jumpers (with and without digging activity);
(6) cursorial forms that also frequently swim. Arboreal

146

A. ELISSAMBURU AND S. F. VIZCAINO

Table 1. List of caviomorph rodents studied, including habits, body mass (in kg) and standard deviation (SD), and number of specimens (n)
Species (common name)

Habits

Body mass

SD

Lagostomus maximus (viscacha)

Myocastor coypo (nutria or coypu)
Galea musteloides (yellow toothed cavy or cuis)
Microcavia sp. (mountain cavy)
Cavia aperea (cavy)
Dasyprocta sp. (agouti)
Dolichotis patagonum (mara)
Hydrochaeris hydrochaeris (capybara)
Lagidium viscacia (mountain viscacha)
Chinchilla sp. (chinchilla)

Digger
Occasional digger
Occasional digger
Occasional digger
Generalised
Cursorial
Cursorial
Cursorial-swimmer
Jumper
Jumper

5.01
3.73
0.4
0.38
1.38
2.97
9.18
48.95
2.5
0.59

1.63
0.67
0.21
0.11
1.02
0.1
1.76
12.16
0.44
0.09

5
4
2
11
9
3
6
2
3
5

forms (e.g. Coendu, Erethizon), whose adaptations are

constrained by different stresses, merit their own study.
Cursorial forms include the dasyproctid Dasyprocta
(agoutis) and the cavid Dolichotis patagonum (Patagonian
hare). Of these, D. patagonum is highly adapted to
cursorial life and is the only caviomorph rodent with
well-marked cursorial adaptations, such as a high degree
of limb extension (Macdonald, 1995: 694). It commonly
takes refuge in vegetation and can occupy empty burrows
and build simple burrows for breeding (Nowak, 1999).
Dasyprocta often builds burrows in riverbanks or under
the roots of trees (Nowak, 1999) and caches excess food in
shallow pits (Smythe, 1978; Emmons, 1990). It develops
appreciable speed and resembles small ungulates in its
locomotor behaviour (Gambaryan, 1974; Grand, 1978
cited in Biknevicius, 1993).
Generalized forms are represented by the cavy Cavia
aperea, which shelter in brush piles and clumps of
vegetation and do not dig burrows (Rood, 1972).
Occasional digger forms include the cavids Microcavia
(mountain cavies) and Galea (yellow-toothed cavies or
cuis), and the myocastorid Myocastor coypus (nutria or
coypu). These taxa dig burrows and also make use of
the abandoned burrows of large mammals (Rood, 1972;
Nowak, 1999). Myocastor coypus has semi-aquatic habits,
using its limbs to swim, rather than lateral undulations of
its vertebral column, with the head and back above the
water, and the tail straight. Elongated hindlimbs and partially webbed digits provide propulsion (Pugliese, 1983).
Digger forms are represented by the chinchillid
Lagostomus maximus (viscacha), which uses the forelimbs
chiefly for digging long burrow systems. It can escape with
speeds of 40 km/h and alternates running with jumps of
3 m and swift turns (Nowak, 1999).
The cursorial-swimming forms are represented by the
hydrochaerid Hydrochaeris hydrochaeris (capybara, body
mass 2779 kg). It resembles Cavia, but is much larger
and has a proportionally shorter body. The limbs are
short. It does not dig (Cabrera & Yepes, 1960), but
runs in an ungulate-like fashion and is a good swimmer
(Nowak, 1999). The limbs, adapted for both running and
swimming, are very short and have a thick interdigital
membrane that favours locomotion in water (Cabal, 1983).
Water is used primarily as a place of refuge and most
normal activity is on land (Schaller, 1976).
The jumping forms include the chinchillids Lagidium
viscacia (mountain viscacha, body mass up to 3 kg) and

Chinchilla spp. (chinchilla, female body mass 0.8 kg and

male body mass 0.5 kg), which run and jump on rocks with
great agility. Lagidium viscacia is a poor digger and is only
rarely found in caves. Chinchillas shelter in crevices and
holes among rocks (Nowak, 1999).
Much work has been devoted to the general structure
and function of the legs in mammals (e.g. Howell,
1944; Gray, 1968; Farish & Jenkins, 1971; Alexander,
1981, 1985, 1989; Hildebrand, 1985; Biewener, 1990;
Raich & Casinos, 1991; Carrano, 1997; Christiansen,
1999; Fischer et al., 2002; Witte et al., 2002 and
references therein). Recent contributions have also
emphasized the morphofunctional study of particular
clades such as xenarthrans (Vizcano, Farina & Mazzeta,
1999, Vizcano & Milne, 2002), carnivores (Jenkins &
Camazine, 1977; Van Valkenburgh, 1987; Iwaniuk,
Pellis & Whishaw, 1999), ungulates (Kappelman, 1988;
Scott, 1990; Thomason, 1991; Alexander & Pond, 1992),
and rodents (Lessa & Patton, 1989; Lessa & Stein,
1992). For caviomorph rodents, research has focused
on the study of specific functional problems or clades
(Biknevicius, 1993, 1999; Biknevicius, McFarlane &
MacPhee, 1993; Casinos et al., 1993, Casinos, Bodini &
Renous, 1996; Fernandez, Vassallo & Zarate, 2000;
among others), but few studies have attempted to explain
partially overlapping functions (Lessa, 1990) or the
functional relationship between fore and hind limbs.
The purpose of this study was to test the relationship
between function and morphology of proximal and middle
skeletal segments of the limbs in terrestrial caviomorph
rodents based on morphometric and biomechanical analyses.
MATERIALS AND METHODS

Fifty-four specimens of caviomorph rodents belonging to

10 species (Appendix) from collections housed in different
museums of Argentina were studied. All specimens
examined were adults, with the exception of the capybara
H. hydrochaeris.
Fifteen measurements of the long bones (Fig. 1,
Appendix), corresponding to diameters and functional
lengths (lengths among articular surfaces) of the bones
and to muscular insertion sites, were taken with digital
callipers to the nearest 0.01 mm. Some are based on the
previous works of Lessa & Stein (1992), Biknevicius
(1993, 1999), Vizcano et al. (1999), Fernandez et al.

Limb proportions and adaptations in caviomorph rodents

147

DMT

OL

PT
DLH
HL

UL
APDH

TDH

TDU

FL

TDF

TL

TDT

DEH

Fig. 1. Measurements of the long bones. HL, functional humerus length; DLH, deltoid length of the humerus; TDH, transverse diameter
of the humerus; APDH, anteroposterior diameter of the humerus; DEH, diameter of the epicondyles; UL, total ulna length; OL, olecranon
length; TDU, transverse diameter of the ulna; FL, functional femur length; DMT, distal extension of greater trochanter; TDF, transverse
diameter of the femur; TL, tibia length; PT, proximal tibial length; TDT, transverse diameter of the tibia.

(2000) and Vizcano & Milne (2002), but modified to

accommodate the peculiarities of the species studied.
Others were specifically developed for this study.

Functional humerus length (HL) is the distance from

the proximal point of the articular head to the distal
mid-point of the trochlea.
Deltoid length of the humerus (DLH) is the distance
from the proximal point of the articular head to the
distal end of the deltoid tuberosity.
Transverse diameter of the humerus (TDH) is the transverse width of the humerus at the diaphyseal midpoint.
Anteroposterior diameter of the humerus (APDH) is
the anteroposterior depth of the humerus taken at a
distance of 35% of the humeral length from the distal
articular surface (Bicknevicius et al., 1993).
Diameter of the epicondyles (DEH) is the transverse
width of the humerus across the epicondyles.
Total ulna length (UL) is the distance from the tip of
the olecranon process to the distal articular surface
of the ulna.
Olecranon length (OL) is the distance along the long
axis of the ulna from the tip of the olecranon process
to the centre of the trochlear notch.
Functional ulna length (FUL) is the difference between
total ulna length (UL) and olecranon length (OL).
Transverse diameter of the ulna (TDU) is the transverse
width of the ulna at the diaphyseal midpoint.
Functional femur length (FL) is the distance from the
proximal point of the femoral head to the distal point
of the femoral condyles.
Distal extension of greater trochanter (DMT) is the
distance from the proximal point of the femoral head
to the distal end of the greater trochanter. DMT is the
functional equivalent of the proximal femoral length
(PF) of Vizcano & Milne (2002).
Transverse diameter of the femur (TDF) is the transverse width of the femur at the diaphyseal midpoint.
Tibia length (TL) is the distance between the proximal
and distal articular surfaces of the tibia.

Proximal tibia length (PT) is the distance from the

proximal articular surface of the tibia to the distal
point of the tibial tuberosity.
Transverse diameter of the tibia (TDT) is the width of
the tibia at the diaphyseal midpoint.
These measurements were combined to create nine
functional indices that represent attributes of bones and
the development and mechanical efficiency of principal
muscles related to limb function. Some of the indices were
taken or modified from the literature (Hildebrand, 1985;
Lessa & Stein, 1992; Vizcano et al., 1999; Vizcano &
Milne, 2002) and others were developed for this study.
They are listed beginning with the indices of the forelimbs,
from the proximal to the distal segment, and ending with
the indices of the hindlimb in the same order.

Shoulder moment index (SMI) is the deltoid length of

the humerus divided by the functional length of the
humerus (DLH/HL 100). This index is an indication of the mechanical advantage of the deltoid and
major pectoral muscles acting across the shoulder
joint.
Humerus robustness index (HRI) is the transverse
diameter of the humerus divided by functional
length of the humerus (TDH/HL 100). It gives an
indication of the robustness of the humerus.
Epicondyle index (EI) is the epicondylar width of
the humerus divided by functional length of the
humerus (DEH/HL 100). It gives an indication of
the relative width available for the origin of the flexor,
pronator, and supinator muscles of the forearm.
It is considered a good indicator of fossoriality
(Hildebrand, 1985; Lessa & Stein, 1992).
Index of fossorial ability (IFA) is the length of the
olecranon process divided by the functional ulna
length (OL/FUL 100). This index gives a measure
of the mechanical advantage of the triceps and dorsoepitrochlearis muscles in elbow extension. It is
also considered a good indicator of fossoriality

A. ELISSAMBURU AND S. F. VIZCAINO

148

Table 2. Genera means (mm), standard deviation (SD), and number of specimens (n) for each index
SD

Genus
Lagostomus

SMI

(n)

SD

HRI

(n)

54.06 2.37 10.88 0.63

(5)
(5)
Myocastor
44.01 2.7 11.08 1.44
(4)
(4)
Galea
45.16 1.71 7.54 1.34
(2)
(2)
Microcavia
37.85 3.87 7.4 0.45
(11)
(11)
Cavia aperea 39.65 3.98 8.31 1.02
(8)
(9)
Dasyprocta
44.79 1.41 7.15 0.88
(3)
(3)
Dolichotis
44.09 6.2
7.54 0.8
(5)
(5)
Hydrochaeris 49.18 0.72 8.57 0.7
(2)
(2)
Lagidium
45.81 1.71 7.89 0.24
(3)
(3)
Chinchilla
42.44 3.03 7.27 1.06
(5)
(5)

SD

EI

(n)

27.05 2.1
(5)
30.06 .32
(4)
17.08 0.71
(2)
19.26 1.51
(11)
19.65 2.48
(9)
19.05 0.71
(3)
17.41 0.31
(5)
26.34 0.48
(2)
20.21 0.6
(3)
19.82 0.77
(5)

SD

IFA

(n)

28.48 2.21
(5)
24.16 2.19
(4)
26.61 2.9
(2)
23.42
(1)
26.11 2.35
(8)
20.12 1.42
(3)
17.43 0.76
(6)
34.66 2.48
(2)
22.43 1.02
(3)
18
1.39
(5)

(Hildebrand, 1985; Vizcano et al., 1999; Fernandez

et al., 2000; Vizcano & Milne, 2002).
Ulna robustness index (URI) is the transverse diameter of the ulna divided by the functional ulna
length (TDU/FUL 100). It gives an indication of
the robustness of the forearm and the relative width
available for the insertion of muscles involved with
pronation and supination of the forearm and flexion
of the manus and digits.
Gluteal index (GI) is the distal extension of greater
trochanter divided by the functional femoral length
(DMT/FL 100). It gives a measure of the mechanical advantage of the gluteus medius muscle and the
velocity of femur extension.
Femur robustness index (FRI) is the transverse diameter
of the femur divided by the functional femoral length
(TDF/FL 100). It gives an idea of capacity to support body mass and to withstand vertical forces associated with velocity increase (Biewener & Taylor,
1986; Demes et al., 1994).
Tibial spine index (TSI) is the proximal tibia length
divided by the length of the tibia (PT/TL 100).
It gives an indication of strength of the leg and
the relative width available for the insertion site of
the gracilis, semitendinosus and semimembranosus
muscles and the foot flexors. It is important in the
flexion of the leg during the first phase of the gait.
Proximal insertion is related to greater speed during
the initial propulsor movement (Elftman, 1929).
Tibia robustness index (TRI) is the transverse
diameter of the tibia divided by the tibial length
(TDT/TL 100). It gives an indication of the
strength of the leg and the relative width available for
the origins of the muscles acting across the ankle.

SD

URI

(n)

6.91

2.41
(5)
6.17 2.37
(4)
4.63 2.82
(2)
4.73
(1)
5.89 .8
(8)
5.71 0.14
(3)
1.12 0.30
(6)
10.72 1.34
(2)
5.11 0.14
(3)
3.59 0.61
(5)

SD

GI

(n)

17.22 2.05
(5)
12.1 2.13
(3)
14.42 0.49
(2)
14
0.87
(11)
14.19 1.34
(8)
12
0.17
(3)
10.38 1.78
(5)
14.68 2.18
(2)
6.13 .92
(3)
8.34 1.47
(5)

FRI

SD

SD

SD

(n)

TSI (n)

TRI (n)

0.35 0.02
(5)
0.46 0.02
(3)
0.38 0.02
(2)
0.35
(1)
0.39 0.05
(6)
0.33 0.01
(3)
0.28 0.1
(5)
0.39 0.02
(2)
0.32 0.06
(3)
0.27 0.01
(5)

7.18 0.01
(5)
6.83 0.005
(4)
5.41 0.003
(2)
6.98
(1)
7.16 0.01
(8)
7
0.002
(3)
6.55 0.003
(6)
8.87 0.006
(2)
6.18 0.003
(3)
5.36 0.006
(5)

11.17 0.01
(5)
11.58 0.01
(4)
10.32 0.002
(2)
9.83 0.01
(11)
12.43 0.02
(9)
11.03 0.005
(3)
10.01 0.004
(5)
11.07 0.002
(2)
9.07 0.002
(3)
9.03 0.007
(5)

Body masses were estimated to evaluate whether the

indices vary with size. Body mass estimations were
calculated as the average of several allometric equations
of the anteroposterior diameter of the humerus and
transverse diameter of the femur (Biknevicius et al., 1993;
Biknevicius, 1999).
Indices were averaged for each genus and were compared through graphic analysis. Based on the correlation
among ulnar proportions and behaviour of different
lineages of mammals (Vizcano et al., 1999), a functional variation series cursorialgeneralizedoccasional
diggingdiggers is recognized. Relationships among
indices and the functional sequence cursorial through
digger were evaluated with Kendalls non-parametric correlations (Legendre & Legendre, 1998). Values for particular jumping and semi-aquatic taxa were compared to
values for the functional sequence categories to determine
which category each taxon most closely resembles.
Principal components analyses (PCA) were conducted
to explore the variation of data in relation to limb function
and to determine the combination of variables (indices
and body mass) that may be important to explain this
variation. Significance was evaluated using the standard
KaiserGuttman ( > 1) criterion and broken stick models
(Legendre & Legendre, 1998). Correlations between
indices and between indices and log-t body mass were
constructed to complete the relationships study.
Functionally important indices were analysed with
discriminant analysis (DA) among the functional categories recognized within the caviomorph rodent group.
RESULTS
Individual indices

For each genus the mean and standard deviation of each

index were calculated; the data are presented in Table 2
and plotted in Figs 2 & 3. Table 3 shows correlations

 SE

SD
56

12

SMI

Mean
30

HRI

26

10

48

24

22

44

40

20
18

36

16

32

14

38

URI

20

IFA

GI

18

10

34

30

14

26

12

22

10

18

14

13.5

Limb proportions and adaptations in caviomorph rodents

12

EI

28

11

52

16

FRI

0.46

12.0

6
4

TSI

0.09

0.40

0.08

0.34

0.07

0.28

0.06

0.22

0.05

0.16

0.04

TRI

10.5
9.0
7.5
Dasyprocia

Cavia

Dolichotis

Galea

Miicrocavia

Lagostomus

Myocastor

Lagidium

Chinchilla Hydrochaeris

Dasyprocia

Cavia

Dolichotis

Galea

Miicrocavia

Lagostomus

Myocastor

Lagidium

Chinchilla Hydrochaeris

Dasyprocia

Cavia

Dolichotis

Galea

Miicrocavia

Lagostomus

Myocastor

Lagidium

Chinchilla Hydrochaeris

Fig. 2. Box plots of each index by genus for caviomorph rodents, showing mean value, standard error (SE) and standard deviation (SD).

149

A. ELISSAMBURU AND S. F. VIZCAINO

150
HRI

EI

IFA

URI

GI

FRI

TSI

TRI

Body mass

Generalized

Cursorial

Occ. digger

Digging

SMI

Fig. 3. Correlations between indices and functional sequence, and between functional sequence and body mass for caviomorph rodents.
Table 3. Correlation among indices and functional sequence,
including number of specimens (n), value of Kendall correlation
( ), Kendalls transformation (Z), and the probability (P). Asterisks
indicate significant correlations
Variable

SMI
HRI
EI
IFA
URI
GI
FRI
TSI
TRI
Body mass

38
39
39
29
29
37
39
25
29
40

0.10
0.28
0.41
0.58
0.37
0.51
0.04
0.24
0.06
0.14

0.85
2.53
3.65
4.44
2.85
4.44
0.33
1.70
0.49
1.30

P
0.406
0.011
<0.001
<0.001
0.004
<0.001
0.742
0.089
0.622
0.193

among indices and the functional sequence and that body

mass is not correlated with the functional sequence.
In the forelimb, the HRI, EI, IFA, and URI are correlated
with the functional sequence, increasing from cursorial to
digging forms. SMI is not correlated with function in this
sequence.
The values for the humeral indices HRI and EI are
variable among the occasional diggers compared to those
of the cursorial and digging forms, mainly owing to
the lower values of Microcavia and Galea and the
higher value of Myocastor. Hydrochaeris has HRI values
between those of Cavia and Lagostomus, and Lagidium
and Chinchilla have values closer to the lower values.
For the EI, Hydrochaeris has values similar to those
of Lagostomus, whereas Lagidium and Chinchilla have
values similar to those of Dasyprocta, Microcavia and
Cavia.
For the ulnar indices IFA and URI, Dolichotis has the
lowest values (clearly different from the other taxa for
URI) and Lagostomus the highest values. The remaining
genera have intermediate values. Hydrochaeris has values
grater than Lagostomus. Chinchilla and Lagidium have
values between those of Dasyprocta and Myocastor for
IFA and lower to that of Dasyprocta for URI.
In the hindlimb, only GI is correlated with the functional
sequence, increasing from cursorial to digging forms.

FRI, TSI, and TRI are not correlated with the functional
sequence.
For GI, Dolichotis has the lowest and Lagostomus the
highest values. The other genera have intermediate values.
Hydrochaeris has values between those of Myocastor and
Lagostomus. Chinchilla and Lagidium have lower values
than Dolichotis.

Principal Components Analyses

PCA was performed on the indices, using data from

32 specimens of caviomorph rodents (one specimen of
Lagostomus, two of Cavia aperea, 10 of Microcavia, one
of Myocastor, two of Dolichotis and two of Chinchilla
were excluded because of incomplete data).
Three eigenvalues are > 1 and explain 74.66% of total
variance among the data. PC1 explained 50% of the
total variance, PC2 explained 14.31% and PC3 explained
10.34%. The percentage of total variance is greater than
that predicted by the broken stick model (63%, P = 10,
1,2,3 ) and hence is considered important for interpretation
of the results.
PC1 shows some separation of taxa into cursorial and
digging functional types with some interesting features
(Fig. 4). The loadings of the variables (which include the
indices and body mass; Table 4) show that PC1 has a
strong association with an increase of humeral robustness
(HRI), development of the epicondyles (EI), olecranon
proportion (IFA), increase of ulnar (URI), femoral (FRI)
and tibial (TRI) robustness, and with distal length increase
of gluteal insertion of the femur (GI). Moderately high
loadings for the increase of the deltoid spine (SMI)
and the tibial tuberosity (TSI) are present. Body mass
has moderately high loadings on this PC. Cursorial
(Dolichotis) and jumping, non-digging (Chinchilla) forms
have lower values on PC1, followed by jumping, lowdigging forms (Lagidium). Digging (Lagostomus) and
cursorial-swiming (Hydrochaeris) forms, with a forceadapted forelimb, have higher values on PC1. Among
them, the less specialized cursorial forms (Dasyprocta)
close to occasional diggers, Galea and Microcavia,

Limb proportions and adaptations in caviomorph rodents

151

Fig. 4. Biplot of PC1 and PC2 values for caviomorph rodents.

Table 4. Factor loadings on principal components. One asterisk and
two asterisks indicate loadings higher than 0.5 and 0.7 respectively
Component
Variable

PC1

SMI
HRI
EI
IFA
URI
GI
FRI
TSI
TRI
Body mass
Expl.Var
Prp.Totl

0.54
0.75
0.77
0.84
0.80
0.73
0.71
0.58
0.75
0.53
5.001
0.500

PC2

PC3

0.57
0.22
0.32
0.17
0.03
0.38
0.51
0.47
0.09
0.55
1.43
0.143

0.08
0.49
0.43
0.12
0.003
0.09
0.17
0.29
0.50
0.46
1.034
0.103

show values close to zero. The generalized C. aperea

and the occasional digger Myocastor show low positive
values, the latter slightly overlapping the lowest values of
Lagostomus.
PC2 shows some separation of taxa by functional
type (Fig. 4). PC2 had a moderately high correlation
with the decrease of the deltoid spine (SMI) and the
increase of femoral robustness (FRI). Body mass has
a moderately high negative loading on this PC. PC2
clearly differentiates between the generalized C. aperea
and the occasional diggers Galea and Microcavia, with
the higher values, from the other caviomorph rodents.
Differentiation among cursorial and digging forms is
not clear. However, values on the PC2 decrease from
occasional diggers (Myocastor) to diggers (Lagostomus)
and cursorial-swimming (Hydrochaeris) forms, as well
as from the cursorial Dasyprocta to the more specialized
cursorial Dolichotis.

PC3 does not show a relation with functional types.

Only Myocastor, with the lowest values on this PC,
shows a clear separation from the remaining taxa. PC3
has a moderately high correlation with increase of tibial
robustness (TRI); and body mass and with decrease of
humeral robustness (HRI).
Correlations

Table 5 summarizes correlations among indices and body

mass. The strongest correlations (P < 0.05) are for the
pairs of indices HRI-EI (0.85), IFA-URI (0.79), GI-FRI
(0.71) and IFA-GI (0.70). Positive correlations, higher
than 0.5, are for the pairs of indices IFA-TRI (0.61),
URI-TRI (0.61), IFA-FRI (0.58), EI-URI (0.57), FRI-TRI
(0.56), FRI-TSI (0.55), HRI-URI (0.54), EI-IFA (0.51),
and SMI-EI (0.50). Combinations of the remaining of
indices have low correlations (< 0.5).
Figure 5 shows the most important relationship between
indices and log-body mass. The main variation with an
increase of body mass is the greater robustness of the
hindlimb (TRI) and the greater deltoid tuberosity (SMI).
EI has a low correlation with body mass.
Discriminant analysis

DA was performed with the indices HRI, EI, IFA, URI, GI,
FRI, TSI and TRI; i.e. those for which the analyses reveal
a meaningful relationship with the cursorial-digging
functional sequence. Discriminant function analysis was
highly significant (Wilks = 0.0452, F (24,32) = 7.357,
P < 0.00001). Data are correctly classified in 100% of
the cases. Among rodents without previous classification,
Hydrochaeris is assigned to the occasional digging and
digging groups, Lagidium to the occasional digging group,
and Chinchilla to the cursorial and the occasional digging
groups.

A. ELISSAMBURU AND S. F. VIZCAINO

152

Table 5. Correlation among indices and body mass. *, Significant at P < 0.05
Correlation

SMI

SMI
HRI
EI
IFA
URI
GI
FRI
TSI
TRI
Body mass

1.00
0.45
0.50
0.40
0.36
0.35
0.11
0.06
0.25
0.48

HRI

0.45
1.00
0.85
0.46
0.54
0.39
0.39
0.43
0.40
0.32

EI

0.50
0.85
1.00
0.51
0.57
0.35
0.34
0.39
0.45
0.40

IFA

URI

0.40
0.46
0.51
1.00
0.79
0.70
0.58
0.42
0.61
0.30

0.36
0.54
0.57
0.79
1.00
0.49
0.45
0.40
0.61
0.29

GI

0.35
0.39
0.35
0.70
0.49
1.00
0.71
0.47
0.41
0.21

56

TRI

FRI

0.11
0.39
0.34
0.58
0.45
0.71
1.00
0.55
0.56
0.21

TSI

0.06
0.43
0.39
0.42
0.40
0.47
0.55
1.00
0.28
0.11

TRI

0.25
0.40
0.45
0.61
0.61
0.41
0.56
0.28
1.00
0.65

Body mass

0.48
0.32
0.40
0.30
0.29
0.21
0.21
0.11
0.65
1.00

SMI

52
8
48
7

44

40
36

5
32
4
0.8

28
0.4

0.0

0.4

0.8

1.2

1.6

0.8
2.0
Body mass

0.4

0.0

0.4

0.8

1.2

1.6

2.0

Fig. 5. Principal correlations between body mass and TRI and SMI for caviomorph rodents.

DISCUSSION
Cursorials to diggers

In this section the individual analysis of each index is

discussed, including the differences in the mean values
and their correlation with body mass, obtained for
those forms recognized as being part of the cursorial
generalizedoccasional diggerdigger sequence.
Forelimb indices

Generally the forelimb indices are correlated with the

functional sequence but not with body mass. Humerus
robustness index (HRI) and the epicondyle index (EI) in
the humerus, and the index of fossorial ability (IFA) and
ulna robustness index (URI) in the ulna, increase from
cursorial to digging forms. On the other hand, shoulder
moment index (SMI) of the humerus is not correlated with
the functional sequence. Only SMI and EI are correlated
with body mass.
SMI: shoulder moment index

The lowest values correspond to the generalized Cavia

aperea and the occasional digger Microcavia. SMI values
increase in the cursorial, digging, and occasional digging
groups, but reach their highest values in the last two groups

(with exception of Microcavia, as mentioned above); this

possibly reflects different muscular adaptations within
the clade. Functional interpretations associated with SMI
are complicated by the fact that muscles with different
functions (deltoid and pectoralis major) insert on the
deltoid tuberosity. The deltoid muscle is both a shoulder
stabilizer and flexor of the arm and the pectoralis major
acts as a limb protractor during locomotion. Fischer et al.
(2002), however, demonstrated the strong contribution of
the scapula, and to a lesser degree the proximal and distal
limb segments, in production of the propulsive movement
in the forelimb in small therian mammals, including
some caviomorphs. The shape and position of the deltoid
tuberosity probably reflect a compromise in response to
a multiplicity of factors, including the force exerted on
substratum, shoulder stabilization, leg support of body
mass, and, to a lesser degree, recuperation speed of step.
Also, muscular arrangements peculiar to some taxa
should be considered. For example, in Cavia and
Dasyprocta, a separate superficial part of the pectoralis
major inserts onto the distal end of the humeral shaft
rather than onto the deltoid tuberosity (Woods, 1972).
Thus, the SMI would not be expected to reflect only
the combined functions of the deltoid and pectoralis, and
therefore might not be an equally reliable indicator of a
particular adaptation for all caviomorphs.
None the less, SMI was identified as a reliable indicator
of digging ability in armadillos (order Xenarthra), a clade
clearly constrained by this function (Vizcano & Milne,

Limb proportions and adaptations in caviomorph rodents

2002). This led us to expect that there would be a positive

correlation between SMI and digging ability in Galea,
Myocastor and Lagostomus, related to flexion of humerus
and stabilization of shoulder for scratch-digger forms
(Hildebrand, 1985).
HRI: humerus robustness index

The correlation of increased humeral robustness from

cursorial to digging forms, combined with non correlation
of body mass with the functional sequence, is probably
related with the necessity of withstanding muscular stress
on the bone, as would be expected for diggers, rather than
inertial forces, as would be expected for cursorial forms.
The occasional digger category includes forms with a
great variation in body mass. Within this group, body mass
correlates positively with HRI (r = 0.93; P < 0.05), and a
size relationship between these forms may be inferred. The
changes in skeletal and muscular arrangements with size
may be related to reduced values of indices in Microcavia
and Galea and to high values in Myocastor.
EI: epicondyle index

An increase of flexor, pronator, and supinator musculature

of the middle and distal forelimb segments and an increase
of force of the manus occur from cursorial to digging
forms. The medial and distal segments make only a minor
contribution to stride (Fischer et al., 2002), and a more
direct involvement with force development for digging
may be inferred. As for HRI, values for the occasional
digger are variable and a positive correlation among body
mass and EI (r = 0.97) within this group is detected. Thus,
skeletal and muscular arrangement on the humerus may be
influenced by the great variation of body mass within the
occasional digging group. The lower values of the index
in Dolichotis reflect its extreme adaptations to cursorial
locomotion. The cursorial (but not highly specialized)
Dasyprocta has values similar to those of the generalized
forms.

153

forms. However, this is the only Cavia species that does

not dig, and these values may reflect retention owing to
phylogenetic constraint.
URI: ulna robustness index

URI increases from cursorial to digger forms. In addition

to the robusticity of the forearm, URI is also related with
the development of many muscles of the forearm and
manus, such as pronators, supinators, and deep digital
flexors. These are related to the major musculoskeletal
modifications of scratch-diggers with increased strength in
flexing the larger digits and the wrist (Hildebrand, 1985).
As with EI and IFA, URI distinguishes among the different
specializations of cursorial forms, with the lowest values
for Dolychotis. The value for Cavia aperea is high and
similar to those of occasional digging forms, as occurs
with IFA.
Hindlimb indices

Only GI is correlated with the functional sequence,

decreasing from cursorial to digging forms. TRI is
strongly correlated with body mass.
GI: gluteal index

The decrease in values of GI (the extension of the greater

trochanter in relation to the femoral length) from digger to
cursorial forms is apparently related to an increase of the
mechanical efficiency of the gluteus medius muscle for
speed. Although femoral position during fast locomotion
in small therian mammals is mainly passively induced
by sagittal spine flexion, the femur is the most propulsive
hindlimb segment during symmetrical gaits (Fischer et al.,
2002) and a relation with limb speed can be supposed.
As with EI, IFA and URI, GI distinguishes among the
different specializations of cursorial forms. As for IFA
and URI in the forelimb, Cavia aperea has a GI value
similar to those of occasional diggers.
FRI: femur robustness index

IFA: index of fossorial ability

IFA, the proportion of the olecranon process in relation to

the ulna length, is correlated with the functional sequence
in relation to an increase in mechanical advantage of
the triceps and dorsoepitrochlearis muscles in elbow
extension and force development for digging (Hildebrand,
1985; Vizcano et al., 1999). The low contribution of the
ulna to stride, and the similar pattern of elbow torque and
work of the triceps muscle during locomotion of small
mammals (Fischer et al., 2002; Witte et al., 2002), support
the functional association of IFA with force development
for digging rather than for speed. With the exception of
Dolichotis, the values obtained are as expected for scratchdigger forms according to Hildebrand (1985). As with EI,
IFA distinguishes among cursorial forms. Cavia aperea
has high values, similar to those of occasional digging

FRI does not correlate with functional sequence or body

mass. A decrease in values from Cavia to the cursorial
forms Dasiprocta and Dolichotis occurs, rather than an
expected increase based on Biknevicius (1993) proposal
that great mean moments of area indicate that the femora
are relatively more rigid, presumably for high-speed
locomotion. Thus, FRI apparently does not reflect vertical
stresses associated with increased velocity; more likely
it reflects other factors such as increased length of the
femur.
TSI: tibia spine index

Values of TSI do not indicate a relationship with function

of limbs or body mass among functional groups. However,

154

A. ELISSAMBURU AND S. F. VIZCAINO

the cursorial forms Dolichotis and Dasyprocta have the

lower values for this index, which may be explained
by the proximal insertion of the semimembranosus and
semitendinosus muscles associated with speed of the
initial phase of step. On the other hand, this index
correlates positively with body mass within generalized
(r = 0.93), occasional diggers (r = 0.89) and digging
(r = 1) forms, and a particular relation to TSI with size
within these groups is inferred.
TRI: tibial robustness index

The high correlation with body mass shows that tibia

robustness is the principal feature of limbs related with
the support of body mass. This reflects the importance
of the tibia in the support and propulsion of body mass
during locomotion.
In addition to the indices discussed above, two others,
brachial index (BI) and crural index (CI), were analysed.
These have been considered as indicative of how well the
fore and hindlimbs are adapted for speed (Howell, 1944;
Fleagle, 1979; Vizcano & Milne, 2002).
BI is the functional ulna length divided by the functional
length of the humerus, and CI is the length of the tibia
divided by the functional femur length. In our rodent
plot, values of these indices are close to 1, as would be
expected among small mammals with a more crouched
limb stance with an c. 1:1 relationship of the segment
lengths (Witte et al., 2002). Within small mammals,
changes of speed and length of gait are directly related
to movement of the more proximal limb segments (the
scapular pivot in the forelimb and the hip joint or sagittal
lumbar spine movements in the hindlimb). Thus, distal
extension of the limbs does not contribute to the length
of the stride. However, among the rodents included in
this study, Dolichotis, with the highest BI and CI values,
merits attention as it is clearly differentiated from the other
rodents. It is the only caviomorph rodent with well-marked
adaptations to cursoriality, with more elongated limbs
and a straighter-angle limb posture, differing from the
usual rodent pattern. A straighter-angle limb posture, as
observed in many small therians, leads to a more elevated
position of the body and thus a higher scapular or elbow
pivot. Thus, an equivalent or even higher contribution to
stride length can be achieved by lower effective angular
movements. Studies on mid-size mammals (Felis, Vulpes,
Procyon, Capra) strongly suggest that the straighter-angle
limb posture of these forms has some influence on their
kinematics (Fischer et al., 2002). For Dolichotis, the more
extended and more distal extension of the limbs compared
with other caviomorph rodents is supposedly related to its
cursorial habits.

Special cases

This section is devoted to discussion of the indices, the

differences among the mean values, and their correlation
with body mass for those forms that do not fit easily into

the functional sequence and are considered special cases,

i.e. the cursorial-swimmers and the jumpers.
Most indices of H. hydrochaeris are similar to those
expected for diggers (EI, IFA, URI) or occasional diggers
(HRI, GI). This species has the highest TRI. Values
of HRI indicate a more robust humerus than in other
cursorial rodents. Casinos, Bodini et al. (1996) suggest
that Hydrochaeris has high peak stresses on the humerus
because the forelimbs are mechanically more active than
the hindlimbs, and state that duty factor and structural
features of the limbs of Hydrochaeris are those expected
for a normal cursorial mammal of its size. The long
olecranon (IFA) may be related to swimming (Vizcano
et al., 1999) because the way this animal swims implies
strong rather than fast extension of the forearm. It is
feasible that other adaptations for force production by
the forelimb, indicated by the development of flexors,
pronator, and supinator muscles (EI, URI), are also related
to swimming. Furthermore, the efficiency of the gluteal
muscles (GI), similar to that of occasional diggers, may
also be an adaptive response of the hindlimb for swimming
as for large body mass. An important factor to consider
is the relatively large body mass of this species, which
is reflected in the high value of TRI. Another probable
consequence of its large size is that Hydrochaeris also
has a more straight-angled stance than most caviomorphs
except for Dolichotis.
The values of some indices (HRI, IG) for Lagidium
and Chinchilla are similar to those that occur in cursorial
caviomorphs, as is expected based on Hildebrands (1985)
assertion that specializations for velocity, as shortening
of muscles moment arms, are similar in cursorial and
jumping forms. On the other hand, indices related to
muscular development of the forearm (EI, IFA, URI) are
similar to those of occasional digging forms, although
Lagidium and Chinchilla are not good diggers.
Principal Components Analyses

PC1 shows significant separation of taxa by cursorialdigger functional types. The position of the jumping
forms adapted to fast movements and the cursorialswimming forms with forelimbs strongly adapted for
force, as well as the low loading of body mass, support
the importance of this PC for interpreting the variation
between speed and force functions within the cursorialdigging sequence. From the cursorial to the digging
extremes, the increase of epicondyle development (EI),
olecranon proportion (IFA), and humeral (HRI) and ulnar
robustness (URI) in the forelimb, and the increase of
the moment arm of gluteal insertion on the femur (GI)
and femoral (FRI) and tibial (TRI) robustness in the
hindlimb are relevant. Thus, the digging forms have
more robust forelimbs, a higher mechanical advantage of
the triceps and dorsoepitrochlearis muscles, and greater
development of flexor, pronator, and supinator muscles
of the forearm and manus than the cursorial forms. The
hindlimbs of diggers have lower indications of speed (GI)
and higher robustness of femoral and tibial portions than

Limb proportions and adaptations in caviomorph rodents

those of cursorial forms. A strong association between

the variation of indicators of force in the medial and
distal segments of the forelimb and the development of
indicator speed in the proximal segment of the hindlimb is
observed. Furthermore, an increase of humeral (SMI) and
tibial (TSI) force production has a secondary relation with
forcespeed variation. The data indicate that Cavia aperea
partially overlaps with the occasional digging forms on
the PC1, and may reflect the generalized condition of this
species.
PC2 reveals some degree of separation among
the functional groups. This separation is particularly
evident for Cavia with respect to the occasional digger
Myocastor and the digging (Lagostomus), cursorialswimming (Hydrochaeris), and cursorial (Dasyprocta
and Dolichotis) taxa, which have similar low scores on
this axis. The interpretation of PC2 is complex and a
direct relationship with the cursorialdigging sequence
is not observed. A relationship for cursorial and digging
functions can be inferred, however, based on the patterns
of SMI and FRI on PC1: a marked increase in deltoid
development (SMI) occurs in diggers (see discussion of
SMI, above) and a decrease of femoral robustness (FRI)
in cursorial forms. Body mass follows the same pattern as
SMI on both PC1 and PC2.
The capybara has the highest values for PC1 and
the lowest values for PC2, which suggests that its limb
bones are very robust and powerful. The jumping forms
Lagidium and Chinchilla have values close to those for
extreme cursorial forms for PC1 and lower values for
PC2, which suggests that limbs are not adapted for force
but for speed.
PC3 does not show a relationship with functional types,
and variation along this axis is related more to differences
within taxa than among taxa.

155

proximal and distal portions of the hindlimb. Similarly, the

low correlations between GI-TSI and GI-TRI suggest that
proximal and medial portions of the hindlimb do not show
the same mechanical relationships with speed or force
variation. TRI and TSI in the tibia have no correlation
with each other.
The main correlations between the limbs are those
between IFA and GI, FRI and TRI, and URI and TRI.
These correlations reveal important relationships between
the mechanical efficiency for force development in the
medial portion of the forelimb and the mechanical
efficiency for speed in the proximal portion of the
hindlimb. Further, a relationship exists between variations
in force and robustness in the middle forelimb segment
and variations in robustness of the proximal and middle
hindlimb segments.
Body mass is principally correlated with TRI and
with SMI, with both increasing with increasing size.
Correlation between body mass and EI is low. EI, however,
is correlated with SMI, and a particular relationship with
body mass for these indices is possible.
Discriminant analysis

Discriminant analysis strongly differentiates the cursorial,

generalized, occasional digging, and digging forms.
The assignment of the swimming-cursorial Hydrochaeris
to the occasional digging and digging group reflects,
as expected, highly specialized limb adaptations for
force development. Among jumping forms, Lagidium is
assigned to the occasional digging group and Chinchilla
to the occasional digging and the cursorial groups. This
might result from the extreme specialization for speed in
the posterior legs and the more generalized foreleg with
low digging activity of these genera.

Correlations
CONCLUSIONS

The main correlations are those involving the common

measures on which each index for the humerus, ulna, and
femur is constructed, and clearly reveal the relationships
among increase of bone robustness, muscle development,
and force efficiency for each of the limb segments.
For the forelimb, the main correlations are among IFA,
URI, and EI, and between URI and HRI. The occurrence
of these correlations can be related to an increase of the
muscular moment arm, an increase of medial and distal
limb mass, and a more powerful distal segment associated
with the increase of force development in digging forms.
The correlation between URI and HRI indicates a clear
relationship between robustness of the proximal and distal
portions of the forelimb from cursorial to digging taxa.
The low correlations between IFA and SMI and HRI and
between URI and SMI suggest the absence of relationships
among the functional specializations of proximal and
medial segments of the forelimb.
The main correlations in the hindlimb are between FRITSI and FRI-TRI, which indicate that, as for the forelimb,
there are clear relationships between robustness of the

This study demonstrates that there is a strong relationship

between limb proportions and function in caviomorph
rodents. In the cursorialgeneralizedoccasional digging
diggers functional sequence, a main trend of increased
force and muscular development in the middle forelimb
segment is indicated by the robustness of the bones and
the greater moment arm of the main muscles used in
force development. The hindlimbs are also clearly related
to functional sequence. Our results reveal a decrease
in the index of speed efficiency of the femur and an
increase of limb robustness from cursorial to digging
forms. Further, an important relationship exists between
mechanical efficiency for force development in the middle
segment of the forelimb and mechanical efficiency for
speed in the proximal segment of the hindlimb, and
between variations among force and robustness in the
middle segment of the forelimb and variation in robustness
of the proximal and middle segments of the hindlimb.
Although overlap of speed and force functions in the
different segments of the limbs is evident, our results

156

A. ELISSAMBURU AND S. F. VIZCAINO

indicate functional specialization for speed in the proximal

and for force in the medial segments, respectively.
When each index is considered independently, those that
reflect forearm (IFA) and femoral (GI) extension are the
ones that best express the relationship between force and
speed variation in the fore and hindlimbs. Hence, these
indices are the most useful single characters reflecting
the adaptations of each limb. In the forelimb, robustness
of the ulna (URI) follows IFA in importance, although
it differentiates poorly between less specialized cursorial
forms and generalized forms. Both epicondylar width
(EI) and robustness of the humerus (HRI) express the
relationship between force and speed variation, but are not
as useful as IFA, GI and URI in distinguishing occasional
diggers from diggers and from cursorial forms. On the
other hand, EI and HRI are related to body mass in
occasional diggers.
Multivariate analyses, however, show that a set of
variables is directly related to forcespeed function, and
reveals a set of variables that best characterizes limb
adaptations. In addition to the already mentioned IFA,
URI and GI, epicondyle development (EI), and humeral
(HRI), femoral (FRI) and tibial robustness (TRI) are the
principal variables related with the variation of force
speed functions.
Distinct patterns for two indices, moment arm of the
deltoid muscle (SMI) and femoral robustness (FRI),
are associated with particular functions. SMI increases
especially in forms specialized for exerting force (i.e.
the diggers) and FRI decreases among cursorial forms
specialized for speed (i.e. Dasyprocta and Dolichotis).
Body mass is correlated mainly with tibia robustness
(TRI) and shoulder moment (SMI), with both increasing
with size.
The terrestrial caviomorph rodents are a variable group
that parallels but does not achieve the extreme adaptations
of other specialized mammals such as running ungulates
and burrowing armadillos. On the one hand, although
the reduction of digits in running caviomorphs is not
as marked as in most artiodactyls and perissodactyls
(maras Dolichotis spp. have four digits in the forelimb
and three in the hind limb, all of them well developed
and touching the ground), cursorial caviomorphs show,
in accordance with specializations for speed present in
canids and felids among carnivores, muscular reduction
in the distal part of limbs and a short olecranon. The most
specialized cursorial caviomorphs, Dolichotis spp., also
have longer distal segments and maintain a less flexed limb
posture than the remaining caviomorphs. A less flexed
limb posture may also be related to weight bearing in the
capybara H. hydrochaeris, a cursorial form with the largest
body mass of the group.
On the other hand, some caviomorphs show convergences with forms specialized for force, with a clear
increase in muscular development and bone robustness
of the forelimb. In particular, diggers have developed
the same muscular groups and trends in moment
arms relationships that occur in specialized outgroup
counterparts such as armadillos, although several skeletal
adaptations, such as development of muscular insertions,

digits and claws, are not as well marked. So, similar trends
to groups more restricted to functional specialization can
be detected within this anatomically variable group of
rodents.

Acknowledgements

Thanks to Darin Croft, Gerry De Iuliis, Richard Farina,

Nick Milne, Manuela Schmidt, and Martin Fischer for
their comments on the manuscript; to Mariano Gordo
Merino and Diego Verzi, Damian Romero, and Olga
Vaccaro for their assistance in the collections of the Museo
de La Plata; Museo Municipal de Mar del Plata and Museo
Argentino de Ciencias Naturales, respectively. Fernando
Momo gave advice on statistics. This is a contribution
to the projects Agencia Nacional de Promocion Cientfica
y Tecnologica, PICT 76348 and Universidad Nacional
de La Plata N336.

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158

APPENDIX. Measurements of the long bones of each specimen of caviomorph rodent

HL, functional humerus length; DLH, deltoid length of the humerus; TDH, transverse diameter of the humerus; APDH, anteroposterior diameter of the humerus; DEH, diameter of the
epicondyles; UL, total ulna length; OL, olecranon length; FUL, functional ulna length; TDU, transverse diameter of the ulna; FL, functional femur length; DMT, distal extension of greater
trochanter; TDF, transverse diameter of the femur; TL, tibia length; PT, proximal tibial length; TDT, transverse diameter of the tibia. MACN, Museo Argentino de Ciencias Naurales
Bernardino Rivadavia (Buenos Aires, Argentina); MLP, Museo de La Plata (La Plata, Argentina); MMCN, Museo Municipal de Ciencias Naturales Lorenzo Scaglia (Mar del Plata,
Argentina).
Specimen

Species

MLP
MLP
MACN
MACN
MLP
MMMdP
MLP
MLP
MLP
MMMdP
MMMdP
MLP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP
MMMdP

8.IX.98.S
27.IV.95.1
48.85
1083
15.V.96.3
1444
3.VIII.972
3.VII.98.2
1089
4103
3295
5.VI.00.9
299
2489
261
301
302
304
306
289
303
3250

Lagostomus maximus
Lagostomus maximus
Lagostomus maximus
Lagostomus maximus
Lagostomus maximus
Lagostomus maximus
Myocastor coypo
Myocastor coypo
Myocastor coypo
Myocastor coypo
Galea musteloides
Galea musteloides
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.
Microcavia sp.

HL

71.4
72.8
70
64.6
57.6
57.9
57.9
58.9
62.52
31.68
27.3
28.76
34.63
27.21
30.98
31.1
32.89
33.26
26.53
33.71
35.15

DLH

TDH

38.3
39.9
38.6
36.5
28.93
24.5
27.3
24.3
28.34
14.69
12
10.87
12.76
11.74
10.64
10.08
14.09
13.55
10.02
13.57
11.11

8
7.8
6.9
7.4
6.45
5.7
5.8
7.6
7.23
2.69
1.8
2.31
2.67
2.14
2.29
2.19
2.2
2.52
1.79
2.64
2.51

DAPH

DEH

9.4
8.4
7.5
8
6.4
8
6.6
6.9
6.76
3.42
2.4
2.32
3.08
2.87
3.26
2.81
3.39
3.08
2.22
3.09
3.03

20.1
18.2
17.5
19.3
15.68
17.6
17.2
17.6
18.91
5.25
4.8
5.75
7.15
5.9
5.57
5.62
6.28
5.85
5.74
6.41
6.27

UL

OL

98.4
95.3
87.5
87.6
79.59
76.9
84.6
89.3
91.66
33.4
28.4

22.2
20.7
19.8
20.8
16
16
14.9
17
18.61
7.44
5.6

34.73

6.59

FUL

76.2
74.6
67.7
66.8
63.59
60.9
69.7
72.3
73.05
25.96
22.8

28.14

TDU

4.9
4.6
4.1
7.4
3.06
3.7
3.1
6.9
3.36
1.72
0.6

1.33

FL

DMT

TDF

87.1

14.3

8.8

101.5
98.4
94.7

14.3
17.8
18.4

11.7
11.4
10.9

77.7
77.8
77.5
81.25
38.88
32.5
33.24
39.17
31.5
37.02
37.31
40.43
39.49
31.17
38.97
39.96

7.7
9.5
11.49
5.47
4.8
4.81
5.93
4.61
5.27
5.6
5.57
4.94
3.92
5.54
5.44

9.4
8.4
8.7
9.9
4.08
3.3
3
4.2
3.74
3.41
3.22
4.02
3.74
2.66
4.12
3.63

LT

PT

TDT

118
130.7
131.1
124.5
118.7

43.1
48.2
46.1
39.8
38.6

7.5
8.3
11.4
8.9
8.7

98.2
92.7
94.6
99.08
44.04
38.4

40.3
44.2
47.21
16.2
15.1

6.6
6.5
7
6.12
2.47
2

48.29

16.77

3.37

A. ELISSAMBURU AND S. F. VIZCAINO

Museum

fn-82-94
5.VI.00.8
585(330)
585(26)
II.VIII.99.54
651,136
525.4
566(315)
29.XIL.00
144
10.VI.98.1
MA-48
MA-16-26
s/n
335
208
207
1080
206
43.43
31.18
534
44.25
29.XII.00.3
24.VIII.98.4
13037
26,116
39,390
11.VIII.99.41

Microcavia sp.
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Cavia aperea
Dasyprocta sp.
Dasyprocta sp.
Dasyprocta sp.
Dolichotis patagonum
Dolichotis patagonum
Dolichotis patagonum
Dolichotis patagonum
Dolichotis patagonum
Dolichotis patagonum
Hydrochaeris hydrochaeris
Hydrochaeris hydrochaeris
Lagidium viscacia
Lagidium viscacia
Lagidium viscacia
Chinchilla sp.
Chinchilla sp.
Chinchilla sp.
Chinchilla sp.
Chinchilla sp.

27.11
39.3
25.3
21.4
38.3
37.6
44.1
41.4
40
35.49
70.3
65.64
62.19
107.92

10.47
17.7
9.3
14.7
16
19
15.7
16.3
11.62
32.1
29.9
26.84
49.5

103.6
111.4
104.3
105
160
142.4
55.6
58.9
60.3
36.2
35.1
32.1
35
33.9

52.1
38.3
49.9
44.2
79.5
69.3
26.3
27.2
26.5
15
15.3
13
16.5
13.4

1.99
2.5
2
1.8
3.2
3.9
3.8
3.4
3.4
2.87
4.5
4.55
5.05
7.67
7.3
7.7
9.2
8.2
14.5
11.5
4.4
4.5
4.9
2.2
2.7
2.6
2.9
2.1

2.43
4.5
2.5
2.3
4.5
6.9
6.3
6.5
4.2
3.4
5.9
6.13
6.65
10.87

4.96
6.4
5.7
5.1
6.7
8.1
8.2
8.1
7.4
6.57
12.9
12.48
12.3
18.94

10.9
11.7
8.6
9.9
23.3
20.1
5.7
5.6
6.5
2.7
3.6
2.7
3.5
2.7

18.3
18.8
18.3
18.3
41.6
38
11.6
11.6
12.1
6.8
7
6.7
70.1
6.6

43.2
26.1
21.9
40.7
42.7
49.3
45.8
44.1

9.1
6
4.2
7.9
9.6
9.9
9.6
8.5

34.1
20.1
17.7
32.8
33.1
39.4
36.2
35.6

2.1
1
1
2.1
2.1
2
1.9
2.6

76.7
70.7
71.66
185.68
170.45
167.4
193
167
179
163.7
148.2
65
70.5
70.3
49.7
50.7
45
51.5
49.7

13.7
11.53
11.51
26.55
26.11
25.8
28.1
25.5
25.4
43.7
36.7
11.8
12.5
13.4
7.4
8.6
6.8
7.5
7.3

63
59.17
60.15
159.13
144.34
141.6
164.9
141.5
153.6
120
111.5
53.2
58
56.9
42.3
42.1
38.2
44
42.4

3.5
3.45
3.46
1.09
1.54
1.6
1.5
2.1
2.2
14
10.9
2.7
2.9
3
1.1
1.8
1.4
1.6
1.6

33
48
29.5
24.8
47.7
45.8
53.4
50.4
50.1
44.62
82.9
77.95
76.4
129.76

4.57
7.2
4.1
6.7
5.5
7.5
8.2
6.5
6.81
10
9.46
9.02
16.84

3.67
5.7
3.7
2.8
5.6
7.3
7.4
6.8
6
4.1
9.5
8.71
7.98
12.85

122.9
128.9
116.2
122.4
198.5
178.9
83.9
88.7
90.8
55
56.7
51.5
56.9
52.9

13.8
11.2
11.8
11.6
32.2
23.5
5.6
5.9
4.6
5.6
4.9
3.3
4.2
4.8

12.5
12.7
12.3
11.7
22.2
19.6
7.4
8.1
8.4
4.8
4.8
5.2
4.8
5

50.6
31.4
27.6
49.7
48.2
58.5
52.6
50.5
96.1
89.29
89.47
171.33
164.63
163.2
186
167
156.2
173.8
162.5
105.9
112.5
111.2
70.8
68.9
62
69.6
67.6

19.8
9.9
9.7
18.4

25
21.1
32.2
31.12
28.77
76.93
40.95
36.4
42.7
36.1
70.1
59.9
42.3
31.3
33.2
19.8
17.5
16.4
20.3
19.2

3.3
2.3
1.8
3.1
4.1
4.6
3.9
3.5
6.5
6.38
6.35
10.55
10.68
10.8
12.5
11.6
10
16.2
13.7
6.2
6.9
7.3
3.4
3.5
4
3.8
3.4

Limb proportions and adaptations in caviomorph rodents

MMMdP
MLP
MLP
MLP
MLP
MLP
MLP
MLP
MLP
MMMdP
MLP
MMMdP
MMMdP
MMMdP
MMMdP
MLP
MLP
MLP
MLP
MACN
MACN
MACN
MACN
MLP
MLP
MACN
MACN
MACN
MLP

159