O

Social grouping (adults)

Mostly tree holes1-2-6-8-12"14 Female sites have better quality than male sites1 Occasionally: leaf nests or dense foliage8-14

Females form

(matrilineal)3-5-16 groups, males mostly stay alone1-3-5-8-14

Gallery forest:

vegetation tangles Spiny forest: tree holes10

Group sleeping observed, composition unknown10

Bark, nests, branches, vines, tree holes' Leaf nests15

Mostly solitary sleeping' Regular (47% of days) sleeping groups of variable composition15

(A) Tree holes, branches, Mostly tree holes, but also vines, and others2 (B) Mostly nests and leaves1'

Mixed-sex sleeping groups2-4-1'

leaf nests, dense foliage8-9-11, palm leaf cavities8, mostly nests, some tree holes1'

Solitary sleeping

(nonreproductive season). partly unisexual sleeping groups (reproductive

Stability of Long-term stability, sleeping changes by deaths, births, groups or group splitting3-5

No long-term stability

Long-term stability, changes by demographic events4

Stable in composition over several months1'

1 Radespiel et al., 1998; 2 Radespiel et al., 2003a;3 Radespiel et al., 2001b; 4 Weidt et al., 2004;5 Lutermann, 2006; 6 Schmid, 1998; 7Schwab, 2000; 8 Martin, 1972; 9Wright and Martin, 1995; 10Rasoazanabary, 2004; 11 Ratsirarson and Ranaivonasy, 2002; 12Petter, 1962; 13Martin, 1973; 14 Pages-Feuillade, 1988;

15 Dammhahn and Kappeler, 2005; 16Wimmer et al., 2002;17Randrianambinina, 2001; 18Weidt, 2001.

Table 4. Socioecological traits of the most studied mouse lemur species (A: Ampijoroa JBA, B: Ampijoroa JBB, K: Kirindy forest, M: Mantadia National Park)

Trait M. murinus M. berthae M. ravelobensis M. rufus group

Size of home ranges (mean) Males

Females

Overlap of home ranges Male-male Female-female Male-female Smallest social unit

Kinship structure Dispersal regime

Category: social organization Female mate choice

(FMC) Male scramble competition (MSC)

Male contest competition (MCC)

(A) 2.2-5.0 ha, seasonally varying12, (A) 0.7-3.0 ha, seasonally varying2, (A) 3.2 ha5, (K) 2.8 ha, seasonally varying4 (A) 1.2-2.6 ha12, (A) 1.7 ha5, (K) 0.7 lu!

High, many1-16

High1-3'4'6, selective1-6, or many3-4 High, many1-5

Female sleeping group1'6 ', mixed sleeping groups8

Matrilinear female groups6 '

Female philopatry, male dispersal3 ''9-1113

Individualized neighbourhood1

Yes: rejection of mates114-22'24, selective escape22-23

Yes: search strategies1-4-28, no spatial monopolization1^1'18

Yes: temporary mate guarding414, male-male aggression2-4'2114, spatial monopolization8

(K) 5.8 ha27, (K) 4.9 ha29 (A) 3.2-7.9 ha26 (B) 0.4-0.8 ha25

(K) 1.2 ha27, (K) 2.5 ha29 (A) 1.8-2.7 ha26 (B) 0.5-0.6 ha25

High, many29 Moderate, some29 High29

Solitary forager2'-29

No matrilines29 Female philopatry, male dispersal29 Individualized neighbourhood29

Yes: male roaming behaviour2'-29, no spatial monopolization29

High26, many25 High26, many25 High26, many25 Mixed-sex sleeping groups25-26

Under study Under study

Individualized neighbourhood25

High30-31 High30-31 High30-31 Solitary forager vs. unisexual male and female sleeping groups30

Possibly male dispersal31 Individualized neighbourhood

Yes: (A) search strategies26, (A+B) no spatial monopolization25-26

Yes: no spatial monopolization3

( Continued)

Table 4. Socioecological traits of the most studied mouse lemur species (A: Ampijoroa JBA, B: Ampijoroa JBB, K: Kirindy forest, M: Mantadia National Park)—Cont'd.

Trait

M. murin us

M. berthae

M. ravelobensis

M. ritfits group

Sperm competition (SC)

Modes of sexual selection Category: mating system

Yes: large testes15-1', multiple mating414'20'22, multiple paternity4-20-22, mating plugs4 FMC+MSC+ MCC+SC Dispersed multimale/multifemale system19

Yes: large testes1'-2''29, mating plugs2'

MSC+SC

Dispersed multimale/ multifemale system2'

Yes: large testes15-1'-26 MSC+SC

Dispersed multimale/ multifemale system25-26

Yes: large testes32

MSC+SC Dispersed multimale / multifemale system

to to to

1 Radespiel, 2000; 2Schmelting, 2000; 3Eberle and Kappeler, 2002b; 4Eberle and Kappeler, 2004b; 5 Pages-Feuillade, 1988; »Lutermann, 2001; 7Radespiel et al., 2001b; »Martin, 1972; 9Radespiel et al., 2003b; 10Fredsted et al., 2005; nWimmer et al., 2002; 12Radespiel, 1998; 13Fredsted et al., 2004; 14Peters, 1999; 15Schmelting et al., 2000; 16Fietz, 1999; 17Schülke et al., 2004; 18 Radespiel et al., 2001a; 19Müller and Thalmann, 2000; 20 Radespiel et al., 2002; 21 Andres et al., 2003; 22 Eberle and Kappeler, 2004a; 23 Radespiel, unpublished data; 24 Radespiel and Zimmermann, 2003; 25 Weidt et al., 2004; 26 Ehresmann, 2000; 2/ Schwab, 2000; 28 Schwab and Ganzhorn, 2004; 29 Dammhahn and Kappeler, 2005;30 Randrianambinina, 2001;31 Atsalis, 2000;32 Randrianambinina et al., 2003

resting period, others form stable sleeping groups of different composition (see Table 3). Most details are known about the social organization of M. murinus. Modern molecular techniques such as microsatellite analyses revealed that this species forms matrilinear female clusters (Lutermann, 2001; Radespiel et al., 2001b) that are characterized by preferential kin-biased space sharing and communal infant rearing (Lutermann, 2001; Eberle and Kappeler, 2002a).

With regard to the mating system of mouse lemurs all species seem to have a dispersed multimale/multifemale system with a pronounced importance of scramble competition and sperm competition among males (see Table 4 for details and references). In general, spatial monopolization of estrous females does not seem to be possible due to their dispersed nightly activities and reproductive synchrony. Males rather seem to have evolved search and roaming strategies in order to localize potential mates. Detailed behavioral observations are again only available for M. murinus. These have shown that in addition to scramble and sperm competition among males, male contest competition and female mate choice can also influence the reproductive outcome. It therefore appears that this small nocturnal lemur species possesses a complex and variable mating system that allows both sexes to adopt different reproductive tactics (e.g., Radespiel, 2000; Eberle and Kappeler, 2004a,b; Schmelting et al., under review), based on competitive abilities, previous experience, body condition, or receptivity of the partner.

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