Sex and seasonal differences in aggression and steroid secretion in Lemur catta:
Are socially dominant females hormonally ‘masculinized’?
Christine M. Drea
⁎
Department of Biological Anthropology and Anatomy, Duke University, 08 Biological Sciences Bldg., Box 90383, Durham, NC 27708-0383, USA
Received 22 June 2006; revised 12 February 2007; accepted 14 February 2007
Available online 23 February 2007
Abstract
Female social dominance characterizes many strepsirrhine primates endemic to Madagascar, but currently there is no comprehensive
explanation for how or why female lemurs routinely dominate males. Reconstructing the evolutionary pressures that may have shaped female
dominance depends on better understanding the mechanism of inheritance, variation in trait expression, and correlating variables. Indeed, relative
to males, many female lemurs also display delayed puberty, size monomorphism, and ‘masculinized’ external genitalia. As in the spotted hyena
(Crocuta crocuta), a species characterized by extreme masculinization of the female, this array of traits focuses attention on the role of androgens
in female development. Consequently, I examined endocrine profiles and social interaction in the ringtailed lemur (Lemur catta) to search for a
potential source of circulating androgen in adult females and an endocrine correlate of female dominance or its proxy, aggression. I measured
serum androstenedione (A
4
), testosterone (T), and estradiol (E
2
) in reproductively intact, adult lemurs (10 females; 12 males) over four annual
cycles. Whereas T concentrations in males far exceeded those in females, A
4
concentrations were only slightly greater in males than in females. In
both sexes, A
4
and T were positively correlated, implicating the Δ
4
-biosynthetic pathway. Moreover, seasonal changes in reproductive function in
both sexes coincided with seasonal changes in behavior, with A
4
and T in males versus A
4
and E
2
in females increasing during periods marked by
heightened aggression. Therefore, A
4
and/or E
2
may be potentially important steroidal sources in female lemurs that could modulate aggression
and underlie a suite of masculinized features.
© 2007 Elsevier Inc. All rights reserved.
Keywords: Female dominance; Aggression; Masculinization; Androstenedione; Testosterone; Estradiol; Seasonality; Lemur; Spotted hyena
Introduction
Unlike most mammals (Ralls, 1976), many strepsirrhine
primates display a social organization that is characterized by
female dominance over males (Jolly, 1966; Richard, 1987). This
unusual trait has received considerable attention and its
behavioral regulation has engendered significant debate: Some
researchers suggest that males defer to females only in feeding
contexts, via ‘male deference’ (Hrdy, 1981) or ‘female feeding
priority’ (Jolly, 1984), whereas others suggest that females
maintain elevated status through overt aggression against males,
via ‘female dominance’ (Kappeler, 1990a; Pereira et al., 1990).
Regardless of the gradations in intersexual social relationships,
functional explanations for the evolution of this trait, that invoke
benefits to the female, are often linked to reproductive
energetics, maternal investment, or nutritional intake (Jolly,
1984; Richard and Nicoll, 1987; Tilden and Oftedal, 1995;
Young et al., 1990); nevertheless, the proximate mechanism
remains a mystery.
Insight into a potential mechanism of female social
dominance may derive from other unusual features of
strepsirrhines, including sexual size monomorphism (Kappe-
ler, 1990b), absence of bimaturation (Leigh and Terranova,
1998), and ambiguous (Hill, 1953; Ioannou, 1971; Petter-
Rousseaux, 1964) or moderately ‘masculinized’ (Drea and
Weil, submitted for publication) external genitalia. This array
of male-like behavioral, physiological, and morphological
traits calls attention to the possible role of androgens in female
lemur development. Although female animals display aggres-
sion in a variety of social contexts, the neuroendocrine
mechanisms that govern female competitive behavior remain
Hormones and Behavior 51 (2007) 555 – 567
www.elsevier.com/locate/yhbeh
⁎
Fax: +1 919 660 7348.
E-mail address: cdrea@duke.edu.
0018-506X/$ - see front matter © 2007 Elsevier Inc. All rights reserved.
doi:10.1016/j.yhbeh.2007.02.006