ARTICLE IN PRESS

www.elsevier.de/mambio

ORIGINAL INVESTIGATION
Effects of group size on nest attendance in the communally breeding
colonial tuco-tuco
Graciela Izquierdoa, Eileen A. Laceyb,
a
Sección Etologı´a, Instituto de Biologı´a, Facultad de Ciencias. Iguá 4225, Montevideo 11400, Uruguay
b
Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, CA 94720, USA

Received 30 May 2007; accepted 27 September 2007

Abstract
Communal nesting is generally assumed to be adaptive, meaning that it confers a fitness advantage on the
individuals that share a nest site. This advantage may accrue directly to adults, or it may affect adult fitness through
gains in offspring survival. In particular, survival of juveniles reared in communal groups may be greater because
adults are present in the nest more often to provide care to young. To test the hypothesis that communal nesting is
associated with increased adult presence in the nest, we used radiotelemetry to examine patterns of adult nest
attendance as a function of group size for free-living colonial tuco-tucos (Ctenomys sociabilis). Burrow systems of this
social, subterranean rodent are inhabited by 1–6 adult females and, in some cases, a single adult male. Data obtained
from residents of 26 burrow systems monitored during 1996–2000 indicated that the percentage of time that the nest
was unattended (no adult present) did not vary predictably with date or time of day during the period between the
birth and weaning of young. The percentage of time that the nest was unattended, however, decreased significantly as
the number of adults per burrow system increased. This difference was most evident when the percentage of time that
the nest was unattended was compared for lone females versus multi-adult groups. We suggest that increased nest
attendance has important implications for the survival of juveniles reared in multi-adult burrow systems but that this
effect may be confounded by the fitness consequences of other costs and benefits associated with communal nesting in
this species.
r 2007 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.

Keywords: Ctenomys sociabilis; Communal nesting; Group size; Nest attendance; Tuco-tucos

Introduction Macdonald 1981; Pilastro 1992; Ebensperger and

Communal nesting is expected to occur when the
fitness benefits of sharing a nest exceed those of living
and rearing young alone (Gittleman 1985; Lewis and
Pusey 1997; Hayes 2000). The selective factors favoring
communal nesting may act directly upon the adults
that share a nest (e.g., increased predator detection;
Corresponding author.
E-mail address: ealacey@berkeley.edu (E.A. Lacey).
Wallen 2002) or they may act upon the young reared
in communal nests (e.g., increased protection from
infanticide; Brown and Burt 2004; Manning et al. 1995;
Ebensperger 1998), thereby enhancing the fitness of
parents that nest together. Because different selective
factors may have distinct or even competing effects on
the actions of individuals, it is important to understand
how the various potential benefits of communal nesting
interact to shape the behavior of animals that live and
rear their young together.

1616-5047/$ - see front matter r 2007 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.mambio.2007.09.007 Mamm. biol. ] (]]]]) ]]]–]]]

Please cite this article as: Izquierdo, G., Lacey, E.A., Effects of group size on nest attendance in the communally breeding colonial tuco-tuco.
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.09.007
 ARTICLE IN PRESS
2 G. Izquierdo, E.A. Lacey / Mamm. biol. ] (]]]]) ]]]–]]]
For juveniles, one potential benefit of communal
nesting is the increased number of adults available to
provide parental care. Among communally nesting
mammals, parental care occurs in a variety of forms,
including provisioning young with food (Gittleman
1985; Baker et al. 1998; Doolan and Macdonald 1999;
Hayes and Solomon 2004), protecting young from
predators (Pilastro 1992; Hoogland 1995), and helping
young to thermoregulate (Trune and Slobodchikoff
1976; Bozinovic et al. 1988; Jacquot and Vessey 1994;
Cutrera et al. 2003). For at least some forms of care,
increasing the number of adults associated with a nest
may increase the benefits accrued by young. For
example, if the presence of an adult in the nest is critical
to thermoregulation by neonates, then the presence of
more adults or the presence of at least one adult for a
greater total proportion of time may enhance the growth
and survival of all young. Thus, patterns of nest
attendance by adults may offer critical insights into
fitness benefits to juveniles associated with communal
nesting.
Studies of the colonial tuco-tuco (Ctenomys sociabilis)
provide an ideal opportunity to explore the adaptive
consequences of communal nesting. Field research on
these subterranean rodents has revealed that nearly 50%
of burrow systems are occupied by multiple adults,
including male–female pairs and multi-female groups
that may or may not include an adult male (Lacey et al.
1997; Lacey and Wieczorek 2004). During the period
when unweaned young are present, all animals in a
burrow system share a single, communal nest site (Lacey
et al. 1997). Although typically all females in single- and
multi-female groups give birth to young, the per capita
number of pups weaned each year is significantly less for
group-living than for lone females (Lacey 2004), raising
intriguing questions regarding the adaptive bases for
communal nesting in this species.
As a first step toward understanding how communal
nesting influences juvenile survival and, hence, parental
fitness, we explored the effects of group size on patterns
of nest attendance by adults. To test the hypothesis that
communal nesting is associated with increased nest
attendance by adults, we quantified the percentage of
time that nests were unattended (no adult present) as a
function of the number of adults per burrow system.
These data provide the first quantitative analyses of
communal nest use by free-living C. sociabilis and yield
critical insights into the potential benefits of sociality in
this species.
Materials and methods
The population of C. sociabilis studied was located on
Estancia Rincon Grande, Provincia Neuquén, Argentina
(401560 S, 711030 W). The study site, which is described in detail
Please cite this article as: Izquierdo, G., Lacey, E.A., Effects of group size on nest attendance in the communally breeding colonial tuco-tuco.
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.09.007
by Lacey and Wieczorek (2003), consisted of a ca. 10-ha area
of open meadow characterized by seasonal grasses, sedges, and
several species of woody shrubs. Each year, the site contained
13–25 burrow systems occupied by C. sociabilis. Typically,
about half of these burrow systems contained a single adult
female, with the remaining burrow systems occupied by a
mean (71 SD) of 3.0 (71.2) females (range ¼ 2–6; Lacey and
Wieczorek 2004). Approximately 44% of burrow systems also
contained an adult male (Lacey and Wieczorek 2004), yielding
a mean (71 SD) population density of 4.0 (72.7) adults per
hectare.
C. sociabilis is almost exclusively subterranean. Individuals
typically emerge only briefly from their burrows to crop
surface-growing vegetation and, while at the surface, the
animals rarely emerge more than half a body length from their
burrows. Members of the study population were captured as
they emerged to forage using hand-held nooses (Lacey et al.
1997). Capture localities were recorded to the nearest meter
using a geo-referenced grid (4 m  4 m cell size) established on
the study site in 1996. The criteria used to determine when all
animals resident in a burrow system had been captured are
described in Lacey et al. (1997). In brief, by placing a small
twig across each entrance to a burrow system, we were able to
determine whether uncaptured animals remained in that
system; if no twigs were displaced during two consecutive
foraging periods (morning and evening), we considered all
animals in that system to have been captured.
Upon first capture, members of the study population were
individually marked by injecting a magnetically coded bead
(IMI-1000 Implantable Transponders, BioMedic Data Sys-
tems, Seaford, DE) beneath the skin at the nape of the neck.
Implanted beads were read using a hand-held scanner (DAS
4004 Pocket Scanner, BioMedic Data Systems). Typically,
adults in the study population were captured during early to
mid-October, which corresponds to the period when females
are giving birth to their single litter of young per year. Adults
were recaptured and juveniles were captured for the first time
between mid-November and mid-December, as soon as young
of the year began foraging for themselves on surface-growing
vegetation (age: ca. 4–5 weeks). Used in conjunction with
radiotelemetry (see below), this protocol allowed us to
determine the number and identities of adults resident in
almost all burrow systems occupied during 1996–2000.
Because the extent of a burrow system cannot be accurately
assessed from surface evidence of activity (e.g., freshly
excavated dirt, active burrow entrances), capture localities
alone do not provide a reliable means of determining which
adults share a burrow system. Consequently, we used radio-
telemetry to confirm that animals captured in close proximity
to one another occupied the same burrow system. Adults
captured in October were fitted with small (o7 g) radiocollars
consisting of an acrylic-encased transmitter (SM1-Mouse
transmitters, AVM Instruments, Inc., Colfax, CA) attached
to a plastic cable tie. Following their release, the locations of
radiocollared animals were determined several times per day
using hand-held antennas and receivers (Yagi antennas, CE-12
receivers, AVM Instruments, Inc.). Previous research has
revealed that localities recorded using this procedure are
accurate to within 0.5 m (Lacey et al. 1997). Individuals that
exhibited extensive (466%) spatial overlap and that shared
the same nest site were considered resident in the same burrow
 ARTICLE IN PRESS
G. Izquierdo, E.A. Lacey / Mamm. biol. ] (]]]]) ]]]–]]]
system (Lacey et al. 1997). Radiocollars were removed when
adults were recaptured during November and December.
Radiotelemetry was also used to document patterns of nest
attendance by adult C. sociabilis during the period between the
birth and weaning of young. Previous field studies (Lacey et al.
1997) have revealed that during this portion of the reproduc-
tive cycle, all residents of a burrow system share a single,
communal nest. For each burrow system monitored during
this study, the location of the nest was identified by
determining the location at which all radiocollared adults
spent the majority of their time (Lacey et al. 1997). This was
done by recording the locations of all radiocollared animals
X6 times per day (minimum of 1 h between successive data
points) for a period of 3–4 days. In addition, the locations
of radiocollared animals were determined after sunset.
C. sociabilis is strictly diurnal and adults only rarely leave
the nest during non-daylight hours (Lacey et al. 1997). As a
result, telemetry data obtained at night provided a convenient
means of determining nest locations.
Once the nest site for a burrow system had been determined,
the presence of adults in the nest was monitored by placing a
Yagi antenna directly over the nest site, with the long axis of
the antennae held perpendicular to the soil surface. The
identities of all adults present in the nest were determined by
quickly scanning through the radiotransmitter frequencies for
adults resident in that burrow system and noting the intensity
of associated radio signals. Intensity was assessed both aurally
and by noting the amplitude of the signal as indicated on the
radio receiver. This procedure provided an unambiguous
means of determining whether an individual was present in
the nest and produced no detectable effects on adult behavior.
Scans of adult presence were repeated at 5-min intervals for a
total of 30 min. Typically, up to three 30-min scan sessions
were completed daily between 0800 and 2100 h, with a
minimum of 2 h allowed between scan sessions for the same
burrow system. The times at which scans were completed for a
given burrow system varied so that all portions of the day were
represented in the data collected from each system.
Because the number of adults per burrow system varied
(Lacey et al. 1997; Lacey and Wieczorek 2004), we use the term
‘‘social unit’’ to describe the set of animals – adults and
juveniles – that were resident in a single burrow system during
a given field season. In terms of benefits to juveniles of
communal nesting, the variable that seemed most likely to
influence offspring growth and survival was the percentage of
time that the nest was unattended (no adults in the nest) during
each 30-min scan session. Although tuco-tucos produce
relatively precocial young (Lacey and Ebensperger 2007),
studies of captive animals indicate that juvenile C. sociabilis
remain closely associated with the nest for the first few weeks
following birth (M. Soares pers. comm.). As a result, the
percentage of time that the nest is unattended may reflect
periods during which parental care is not available to young.
Although each social unit represented an independent data
set, data collected from the same social unit during different
30-min scan sessions were not independent and thus time series
analyses were used to assess whether nest attendance was
influenced by the date or the time of day when scans were
conducted. For each social unit, the percentage of time that the
nest was unattended was calculated per scan session and per
day by dividing the number of scans during which no adults
Please cite this article as: Izquierdo, G., Lacey, E.A., Effects of group size on nest attendance in the communally breeding colonial tuco-tuco.
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.09.007
3
were in the nest by the associated total number of scans. Time
series analyses were then run using date and time of day as the
independent variables. t-tests were used to compare correlation
coefficients from all social units against the null hypothesis of
no consistent relationship (mean r ¼ 0) between nest atten-
dance and date or hour of data collection (Zar 1999).
The effects of social unit size on nest attendance were
examined using ANOVAs and linear regressions. For all other
analyses, parametric statistical tests were used whenever the
associated assumptions of normality and homogeneity of
variances were met. All statistical analyses were completed
using Statistica 6.0 (Statsoft, Inc., 2002). Throughout the text,
means are reported 71 SD.
Results
Nest attendance by adult C. sociabilis was monitored
for 61 individuals in 26 social units resident on the study
site during 1996–2000. All adults in 24 (92.3%) of these
social units were monitored via radiotelemetry. Although
one adult in each of the remaining social units was not
captured until after telemetry data had been collected,
visual inspection of the data obtained from these groups
revealed no apparent differences in nest attendance
between these social units and same-sized groups in
which all adults were monitored. As a result, analyses of
nest attendance for the two incomplete social units were
based on the total number of adults resident in each
burrow system. Twelve of the social units monitored
contained a single adult female; the remaining social units
each contained multiple adult females (mean ¼ 2.071.1
females per social unit, range ¼ 2–4; Table 1). Ten
(38.5%) social units contained an adult male; the
distribution of males between burrow systems occupied
by one versus multiple adult females was not significantly
different from that expected if males are equally
represented in both types of social units (w21 ¼ 1:38,
P40.05). For each social unit, a minimum of 10 30-min
nest attendance scans (mean ¼ 24.7710.8 scans per
social unit; range ¼ 10–48) was completed over at least
5 different days (mean ¼ 23.6710.5 days per social unit,
range ¼ 5–40) between the birth and weaning of pups
reared by that social unit.
As expected, within each multi-adult social unit, all
adults shared a common nest site; in no case did the nest
Table 1. Composition of the social units monitored during
this study
Number of adult females Total
1 2 3 4
Male present 3 3 2 2 10
No male present 9 1 5 1 16
Total 12 4 7 3 26
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sites identified for individuals differ among members of

the same social unit. The percentage of time that the nest
was left unattended (no adults present) did not vary
predictably with the date of data collection (mean
r ¼ 0.02470.352, N ¼ 26; t-test: t23 ¼ 0.35, two-
tailed P40.05), providing no evidence that patterns of
adult activity changed predictably as a function of
juvenile age. Although the percentage of time that the
nest was unattended appeared to increase over the
course of a day (mean r ¼ 0.19370.222, N ¼ 26; t-test:
t23 ¼ 4.43, two-tailed Po0.01), a strict interpretation of
this result indicates only that there was a consistently
non-zero relationship between time of day and nest
attendance among the social units monitored. Closer
inspection of these data revealed that, across social
units, both positive and negative relationships occurred
between these variables. Further, significant correlations
between time of day and percentage of time that the nest
was unattended were detected for only 2 (7.7%) of the
26 social units monitored, indicating that, within social
units, nest use by adults was not strongly temporally
patterned. Given these findings, we did not partition the
data according to date or time of day at which scans
were completed. Instead, for each social unit, all scans
were pooled for the following analyses of nest atten-
dance by adults.
Because not all social units in our sample contained
an adult male (Table 1), we examined the effects of the
presence of a male on patterns of nest attendance by
females. A two-way analysis of variance revealed no
significant effect of a male on the percentage of time that
the nest was unattended (ANOVA: F1,18 ¼ 1.23,
P ¼ 0.283). Consequently, we did not distinguish
between the sexes but instead used the total number of
adults resident in a burrow system for subsequent
analyses of the effects of group size on nest attendance.
When the total number adults per burrow system was
considered, there was a significant effect of social unit
size on the percentage of time that no adults were
present in the nest (one-way ANOVA: F4,21 ¼ 12.57,
Po0.001). Visual inspection of the data suggested that
the percentage of time that the nests of lone females
were unattended was considerably greater than that for
social units containing multiple adults (Fig. 1); post hoc
comparisons revealed significant contrasts between data
from lone females and those from all other sizes of social
unit (Tukey’s HSD tests, all Po0.01), suggesting that
the primary determinant of the percentage of time that a
nest is left unattended is the presence of one versus
multiple adults in a burrow system. Accordingly,
although the percentage of time that the nest was
unattended declined significantly as a function of the
number of adults in a burrow system when all social
units were considered (R2 ¼ 0.533, N ¼ 26, F1,25 ¼
27.35, Po0.001; Fig. 1), no significant relationship
between number of adults and percentage of time the
Fig. 1. Percentage of time that nests of C. sociabilis were
unattended (no adult present) as a function of the total number
of adults resident in a burrow system. Data are from 26 social
units monitored during 1996–2000. Burrow systems containing
an adult male are indicated. The regression line indicates a
significant (Po0.001) negative relationship between number of
adults and percentage of time the nest was unattended.
nest was unattended was detected among social units
containing X2 adults (R2 ¼ 0.114, N ¼ 17, F1,15 ¼ 1.94,
P40.100).
Discussion
Our data indicate that nest attendance by free-living
C. sociabilis was influenced by the number of adults that
shared a burrow system. When all social units were
considered, the percentage of time that the communal
nest was unattended (no adults present) decreased
significantly as the number of adults per burrow system
increased. The presence of an adult male appeared to
have no effect on the percentage of time that the nest
was unattended. When the number of adults per social
unit was considered regardless of sex, the only sig-
nificant contrasts detected were between lone females
and social units containing multiple adults. Thus, the
greatest effect of communal nesting on nest attendance
occurred with the addition of a second adult to social
units containing only a single female.
Reducing the time that the nest is unattended may be
beneficial to juveniles in several ways. In general,
because the presence of an adult is essential to many
forms of parental care, increasing the percentage of time
during which at least one adult is present in the nest may
increase the overall availability of such care, potentially
leading to enhanced juvenile growth and survival (Hayes
and Solomon 2006). More specifically, the presence of
one or more adults may have thermoregulatory benefits
for young; juvenile ctenomyids do not thermoregulate
effectively until they are more than 15 days old (Cutrera

Please cite this article as: Izquierdo, G., Lacey, E.A., Effects of group size on nest attendance in the communally breeding colonial tuco-tuco.
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.09.007
 ARTICLE IN PRESS
G. Izquierdo, E.A. Lacey / Mamm. biol. ] (]]]]) ]]]–]]]
et al. 2003; Zenuto et al. 2002) and, hence, regular nest
attendance may reduce the chances of hypothermia of
young. At the same time, the presence of an adult may
increase the tendency for pups to remain inside the nest
(M. Soares, pers. comm.), thereby decreasing the risk of
predation as well as hypothermia. If, as these arguments
suggest, the regular presence in the nest of X1 adult
increases juvenile survival, then decreasing the time that
the nest is unattended may be an adaptive consequence
of group living in C. sociabilis.
If increased nest attendance is an adaptive benefit of
group living, why do individuals in multi-female social
units rear fewer young to weaning than lone females? On
an annual basis, the per capita direct fitness of female
C. sociabilis declines significantly as a function of group
size (Lacey 2004), indicating that social units in which
nest attendance by adults is greater produce fewer
surviving pups per female. Communal nesting is a
complex social phenomenon that likely reflects the
combined effects of multiple selective factors that confer
both benefits and costs on animals that live and rear
their young together (Alexander 1974; Hayes 2000). As a
result, while increasing nest attendance may improve
juvenile survival, other aspects of communal nesting
(e.g., reproductive skew, infanticide, and other manifes-
tations of increased competition among burrow-mates:
Alexander 1974; Clutton-Brock 1998; Ebensperger 1998;
Woodroffe and Macdonald 2000) may counter this
potential fitness gain. Hence, to understand the net
fitness consequences of communal nesting, it is neces-
sary to consider the full range of selective pressures
leading to this behavior. Future studies of communal
nesting in C. sociabilis will address this issue by
exploring other mechanisms (e.g., communal nursing)
by which young may benefit from increased parental
presence, as well as by exploring the fitness conse-
quences to adults of sharing a nest with conspecifics.
Acknowledgements
For permission to work on Estancia Rincon Grande,
we thank Alain Thouyaret and the Delegación Tecnica
de Parques Nacionales Argentinas, in particular Claudio
Chehebar and Eduardo Ramilo. For logistic assistance,
we thank Miguel Christie, Gustavo Iglesias, and the
members of the Sociedad Naturalista Andino Patago-
nica in Bariloche, Argentina. Field assistance was
provided by A. Chiesa, I. Tomasco, and, in particular,
M. Soares and J. Wieczorek. Previous versions of this
manuscript benefited from comments by G. Francescoli,
M. Hauber, M. Soares, and J. Wieczorek. Melitta
Meneghel kindly translated the abstract to German.
Financial support was provided by the Comision
Sectorial de Investigacion Cientifica and Programa de
Desarrollo de las Ciencias Básicas (Uruguay), and the
Please cite this article as: Izquierdo, G., Lacey, E.A., Effects of group size on nest attendance in the communally breeding colonial tuco-tuco.
Mamm. Biol. (2007), doi:10.1016/j.mambio.2007.09.007
5
Museum of Vertebrate Zoology and Department of
Integrative Biology at the University of California,
Berkeley. This work was conducted in compliance with
all relevant institutional, national, and international
regulations.
Zusammenfassung
Einfluss der Gruppengröße auf die Nestanwesenheit ko-
lonialer Tucotucos (Ctenomys sociabilis)
Gemeinschaftliche Jungenaufzucht wird als adaptiv
angenommen, wenn es den Individuen, die dasselbe Nest
benutzen, einen Fitnessvorteil verleiht. Dieser Vorteil
kann direkt für die adulten Tiere nützlich sein oder deren
Fitness durch längere Überlebensrate ihrer Nachkommen
beeinflussen. Insbesondere Jungtiere, die in gemeinschaf-
tlichen Gruppen aufgezogen wurden, dürften eine längere
Überlebensrate durch die häufigere Anwesenheit meh-
rerer pflegeversorgenden Adulten haben.
Es wurden radiotelemetrische Analysen durchgeführt,
um zu prüfen, wie gemeinschaftliche Vermehrung mit
erhöhter Anwesenheit adulter Tiere in Verbindung steht,
um damit die Hypothese zu testen, dass das Anwesen-
heitsmuster adulter Tiere im Nest abhängig von der
Gruppengröße freilebender kolonialer Tucotucos (Cte-
nomys sociabilis) ist. Die Bausysteme dieses sozialen,
unterirdischen Nagetieres werden von 1–6 adulten
Weibchen und in einigen Fällen von einem einzigen
adulten Männchen bewohnt. Die Daten, die wir von
1996 bis 2000 von 26 überwachten Bausystemen
erhielten, deuteten an, dass die Zeit, in der das Nest
ohne adulte Tiere zwischen Geburt und Absetzen war,
nicht von Datum oder Tageszeit abhing. Die Zeit, in der
das Nest unbeaufsichtigt war, verminderte sich mit der
Zunahme an erwachsenen Tieren für jedes Bausystem
erheblich. Als die Zeit, in der das Nest unbegleitet war,
für solitäre Weibchen mit der von Gruppen mehrerer
Weibchen verglichen wurde, war der Unterschied
offensichtlich. Es wird vorgeschlagen, dass zunehmende
Nestanwesenheit eine wichtige Rolle für das Überleben
von Jungtieren, die in ‘‘Multi-Erwachsenen’’-Bausyste-
men aufgezogen werden, spielt. Dieses könnte jedoch
mit den Fitnessfolgen anderer Kosten und Vorteile, die
mit der gemeinsamen Jungenaufzucht in dieser Art
verbunden sind, verwechselt werden.
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