Primates exhibit a diversity of locomotion types from quadrupedalism to leaping (Napier and Walker, 1967; Hunt et al., 1996). A familiar locomotor category in primatology is vertical clinging and leaping (VCL). The main primate groups that exhibit this locomotion are tarsiers and some strepsirrhines. Tarsiers, indriids, sifakas, some galagos and calltrichids are morphological specialized for this specific locomotion (Napier and Walker; 1967; Terranova and Dagosto, 1994).Ashton and Oxnard (1964) were the first to attempt classifying primate locomotion based upon behavioral classifications.
Prost (1965) was important for his consideration of the difference between positional behaviors and locomotive behaviors. Napier and Walker (1967) argued for a new category type of prosimian (strepsirrhines and tarsiers) locomotion called vertical clinging and leaping. Vertical clinging and leaping involves the adopting of an orthograde (upright) posture at rest on a vertically oriented support that allows primates to initiate movement through extending their hind limbs. They then leap from one support to another whilst rotating their bodies to land on another vertical support (Napier and Walker, 1967; Stern and Oxnard, 1973; Demes et al., 1991; Crompton et al., 1993; Fleagle, 2013).
Most arboreal primates have been shown to exhibit vertical clinging (Dagosto, 1994; Fontain, 1990; Garber, 1991; Napier and Walker, 1967). Primates who most often apply vertical clinging and leaping are large bodied strepsirrhines, such as indris and sifakas, to small body primates, including galagos and tarsiers (Terranova and Dagosto, 1994) (Figure 1). Propithecus verreauxi (a large bodied lemur ranging from 3700 to 4280 grams) habitually uses vertical clinging and leaping (Napier and Walker, 1967; Smith and Jungers, 1977; Walker, 1979; Johnson, 2012). Propithecus coquereli is another example of large bodied habitual vertical clinging and leaping primate (Figure 2). Hapalemur (bamboo lemurs) also exhibits vertical clinging and leaping but infrequently when compared to other vertical clinging and leaping primates (Gebo, 2011).
Vertical clinging and leaping has also been observed in calltrichids (Napier and Walker, 1967; Crompton and Andau, 1986; Dagosto et al., 2001). Cartmill (1974) suggested that the nails of Callicebus sp. allow them to better move on vertical supports. Pygmy marmosets will often take position vertically on substrates to feed on sap allow them to make the vertical posture part of vertical clinging and leaping (Kinzey et al., 1975). A constraint on locomotion is often diet. For instance, indriids, sportive lemurs and bamboo lemurs have a low energy diet with their focus on leafy food resources. Folivory acts as a dietary constraint on vertical clinging and leaping, thus the morphological adaptions for vertical clinging and leaping differ depending on relative size and diet (Gebo, 2011).
Napier and Walker (1967) argued vertical clinging and leaping to be the earliest locomotion specialization of primates. They suggested several features of vertical clinger and leapers could be related back to the posture/movement of Eocene primates thus the morphology of vertical clingers and leapers were pre-adaptive to the other primate locomotion adaptions (Napier and Walker, 1967). However, this hypothesis of vertical clinging and leaping as an early pre-adaptive trait was quickly challenged by Cartmill (1972), Martin (1972) and Stern and Oxnard (1973).
Vertical clinging and leaping as an ancestral condition would not be possible since there is a lack of generalized morphology shared between haplorrhine and strepsirrhines vertical clingers and leapers. For example, the hind limb anatomies are different between differently sized species that practice vertical clinging and leaping and thus no overall anatomical pattern exists (Stern and Oxnard, 1973). Overall the early locomotion technique was likely generalized (Martin, 1972; Cartmill, 1972). Instead of being an ancestral state of locomotion vertical clinging and leaping likely evolved as several different lineages since they lack an overall anatomical feature and lack a similar feeding environment (Gebo, 2011).
Work Cited/Read more
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female dominance, which is defined as the ability of females to consistently evoke submissive behavior from all males (Jolly, 1966; Richard, 1987; Kappeler, 1993). Female dominance can only occur in contexts of male submission (Hrdy, 1981). When males exhibit spontaneous male submission to females in the absence of female aggression, this is termed deference (Kappeler, 1993a). In the absence of male deferential behavior, females can elicit submissive male behavior through the use of aggression (Sauther, 1993).
Another animal that has shown female dominance is the spotted hyena (Kruuk, 1972; Frank, 1989; Smale et al., 1993). However, they do not exhibit true female dominance, as seen in lemurs, since female hyenas do not consistently dominate males in all contexts (Smale et al., 1993; Frank et al., 1989). Jolly (1966) found most dominance interactions that occur among lemurs occur in the context of feeding and often occur with the female supplanting the males. Female lemurs will have feeding priority after winning aggressive interactions with males who submissively vocalize and retreat (Jolly, 1966, 1967, 1984).
energy and maximize scare resources, and suggested three hypotheses to explain the evolution of female dominance in lemurs. The energy conservation hypothesis emphasizes the harsh island climate that selected for female dominance. The evolutionary disequilibrium hypothesis suggests that lemur traits are the result of a transition from nocturnal lifestyle to a more diurnal lifestyle. The energy frugality hypothesis predicts that certain lemur traits developed for maximization of food resources and thus to conserve energy (Wright, 1999). Wright (1999) suggested that the harsh and unpredictable island environment ofMadagascar led to the evolution of several behavioral characteristics, including female dominance, targeted female to female aggression, high female mortality rate, strict seasonal breeding and weaning synchrony.
Richard (1987) argued that females have a strong interest in maintaining their position of social dominance in order to maintain their priority access to food. Males will accept female dominance in order to further their reproductive interests through promoting the survival of their unborn young. Hrdy (1981) argues that the females’ reproductive success is dependent on the ability to obtain resources, and thus that the social dominance of female lemurs and male deference to females is necessary in order to conserve energy for the brief and annual mating season. Jolly (1984) suggested that female dominance occurred due to seasonal stressors on the females.
Sauther (1993) in her study of ring-tailed lemurs in Beza Mahafaly, Madagascar found that her data supported female dominance as a response to the high reproductive costs experienced by female lemurs in the seasonal and harsh environment of Madagascar. Sauther (1993) suggested that males are both direct and indirect feeding competitors of females, and that this competition often coincides with periods of high-cost reproductive states, including lactation during periods of low food availability.
Female dominace is exhibited by most lemur species and involves the ability for the female to consistently evoke submissive behavior from all males (Jolly, 1966; Richard, 1987; Kappeler, 1993). emale dominance can only occur in contexts of male submission (Hrdy, 1981). When males exhibit spontaneous male submission to females in the absence of female aggression, this is termed deference (Kappeler, 1993a). In the absence of male deferential behavior, females can elicit submissive male behavior through the use of aggression (Sauther, 1993).
Another animal that has shown female dominance is the spotted hyena (Kruuk, 1972; Frank, 1989; Smale et al., 1993). However, they do not exhibit true female dominance, as seen in lemurs, since female hyenas do not consistently dominate males in all contexts (Smale et al., 1993; Frank et al., 1989). Jolly (1966) found most dominance interactions that occur among lemurs occur in the context of feeding and often occur with the female supplanting the males. Female lemurs will have feeding priority after winning aggressive interactions with males who submissively vocalize and retreat (Jolly, 1966, 1967, 1984).Ring-tail lemurs lack sexual dimorphism, both females and males are the same size (Mittermeier et al., 1994; Sussman, 2000).
catta are endemic to southwestern Madagascar (Sussman, 1977). Madagascar is located 800 kilometers from Southeast Africa. It is the fourth largest island in the world (Swindler, 2002). Southern Madagascar is characterized by a rainy season that occurs from November to March (Jolly, 1966). L. catta are mainly found in forests of dry bush, gallery, deciduous, scrub and closed canopy forests (Mittermeier et al., 1994). L. catta are opportunistic omnivores and their diet mainly consists of fruit, leaves, flowers, herbs, other plant parts, and sap (Sauther, 1992; Mittermeier et al., 1994). Sauther (1992) found that 30% to 60% of their diet consisted of feeding on fruit and 30% to 51% of their diet consisted on feeding on leaves and herbs.
catta live in multi male, multi female groups ranging from nine to twenty two animals but on average have groups of fourteen animals (Sauther, 1991; Sussman, 1991). Females are philopatric and males disperse; dispersing males tend to enter new groups with one or two other males from their original groups (Sauther, 1991; Sussman, 1991). Generally one female is the single highest-ranking female. The highest-ranking female is typically the one to initiate group movement in a certain direction (Jolly, 1966; Sauther and Sussman, 1993). Females form a dominance hierarchy in which certain females will out rank other females depending on the matriline to which they belong; however, every female is more dominant than males. These dominance relations are reinforced by agonistic encounters (Jolly, 1966; Taylor and Sussman, 1985; Gould et al., 1999).
Males within these social groups are peripheral; however, there are often one to three central/dominant males. The male hierarchy is often dependent on age (Sussman, 1992). Generally the central males are more fit than other males, while the lower ranking/peripheral males are older, recently transferred males as well as younger males that have yet to transfer (Sussman, 1991; 1992; Gould, 1996; Gould et al., 1999). The higher-ranking males often have more interactions with females and thus are more likely to mate, but lower-ranking males also mate (Sussman, 1991, 1992; Gould, 1994, 1996; Gould et al., 1999).
Females start reproducing when they are two or three years old and generally havean interbirth ratio of 2 to 3 years (Sauther, 1991). Mating activity in a group will normally last for two to three weeks per year (Sauther, 1991), however, each individual female will only be sexually receptive for one to two days per year (Van Horn and Resko, 1977; Sauther, 1991).
A Primatologist in The Making 