The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-Human Primates / Beth Christine Laas
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The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-human Primates / Beth Christine Laas "The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-human Primates" By: Beth Christine Laas Anthropology Senior Thesis Dr. Martin Nickels, Advisor Spring 2001 Abstract Primatologists have been observing interbreeding in non-human primate species since the early 1960s. Often this interspecies breeding produces a mixture of morphological and genetic features in the hybrid offspring. The purpose of this paper is to address the effects this interspecies breeding has on the morphological, fertility and viability of the offspring. This paper will analyze observational data and available species studies on interbreeding between Papio hamadryas and Papio anubis in a natural environment, as well as the mating of a gibbon and siamang and a rhesus macaque and a baboon in captive situations. It will also examine the different explanations for this interbreeding as well as what perpetuates it. Through this data collection this paper will show that the different explanations of how interbreeding is perpetuated are both integral parts of this behavior. Introduction Interbreeding between different non-human primate species has always been of interest to primatologists and geneticists alike. The results of interbreeding are offspring that are intermediate between their parents in terms of morphology, genetic traits and behavior. In this paper I will analyze the data from existing journals and species studies of a number of different primate species, including two species of baboons in the wild, as well as the interbreeding of macaque species and a gibbon and siamang in a captive situation, in order to determine the effects of hybridization on the offspring. When examining hybridization between different species, certain questions arise such as whether there is a closer resemblance to one parent or the other, or if there is a possible advantage to being a hybrid rather than a more "pure" form. There are also two prevalent explanations as to why interbreeding occurs in the wild and how it is perpetuated. http://www.soa.ilstu.edu/anthropology/theses/laas/index.htm (1 of 15) [11/30/2007 1:20:42 PM] The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-human Primates / Beth Christine Laas These two ideas are presented, and their appropriateness will be determined. The purpose of this paper is to evaluate these ideas and, in a broader sense, address the overall effects of interbreeding on nonhuman primates in general. I will be looking at several different hybrid crosses in order to analyze the morphological and genetic features found in the offspring of each. Morphological, genetic and behavioral attributes found in hybrid offspring are often shown to be a mix of both parents’ features (Moore, et. al.: 1999:120). This paper will analyze these crosses and evaluate their results in order to determine which species’ are more successful and why. In addition, I will also discuss any effects interbreeding has on the physical well being of the offspring, whether positive or negative. I expect to find that the hybrid offspring produced through interbreeding in the wild will exhibit a mixture of morphological and genetic traits inherited from each parent in the cross. Literature Review Interbreeding among nonhuman primate species has been well documented in the wild between two baboon species, Papio hamadryas and P. anubis. Explanations have been proposed as to why these two species keep interbreeding. Suggested explanations why these and other primates interbreed are related to their behavioral and morphological attributes. The observation of these two species interbreeding led to study of further inter-species breeding in captivity, such as the gibbon and siamang or the baboon and rhesus monkey. Initially, the physical attributes were studied, but over time behavioral attributes, social structure, genetic traits, and adaptations were studied also. Extensive observation of the hybrid offspring of the two baboon species in the Awash Valley in Ethiopia began in the early 1970s, particularly with Nagel's fieldwork. Nagel believed the primary motivation behind the interbreeding was the abduction of anubis females by hamadryas males (Nagel: 1973). This ties into Nagel's other idea: that the hybrid zone was fixed, rather than fluctuating. Nagel did not see environmental factors as playing a part in the groups interbreeding and felt that fixed ecological barriers kept the hybrid zone strictly demarcated. Phillips-Conroy and Jolly contradicted these hypotheses. While female abduction did occur, this in and of itself was "…not frequent enough to account for such rapid changes." (Phillips-Conroy, Jolly: 1986:346). They also found in their observations that "…the hybrid zone was in flux, not static" and that it was environmental factors, such as anubis' vulnerability to drought, that http://www.soa.ilstu.edu/anthropology/theses/laas/index.htm (2 of 15) [11/30/2007 1:20:42 PM] The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-human Primates / Beth Christine Laas played the primary role in these two species interbreeding (Phillips-Conroy, Jolly: 1986:346). The two groups were not necessarily restricted to one particular ecological area. Phillips-Conroy and Jolly went even beyond disputing Nagel's explanations about this interbreeding. They had their own ideas that individuals with certain traits perpetuated this inter-group breeding. They thought that individuals whose phenotype resembled the recipient population, those who were otherwise more successful at entering "foreign" populations, or those who had a lower probability of breeding in their own population were maintaining this interbreeding between groups (Phillips-Conroy, Jolly: 1991:353). As for other interspecies crosses, there can be no question about the motivation behind them because they occurred in a captive situation. The gibbon and siamang mating as well as the many attempts at intergeneric crosses between Macaca and Papio all happened in zoos and captive situations (Hill: 1970:217). Any motivation behind this interbreeding would most likely come from being held in the same area together, or through human intervention. Nagel and Phillips-Conroy and Jolly did agree on the calculation of morphological hybrid index (MHI). Using seven different external features to find the MHI, Nagel discovered that there were no "pure" groups; rather there were varying degrees of "hamadryas-like" features and "anubis-like" features (Nagel: 1973). Social structure and behavior can often be affected by interbreeding. Certain innate behaviors of one species can often become confused with those of another species in a hybrid offspring. Sugawara studied the social structure of a group of wild baboon hybrids in the same region as Nagel, Phillips-Conroy and Jolly. What Sugawara found was that while the social structure in some of the study groups was closer to that of the anubis type, it did exhibit hamadryas characteristics as well, and vice versa (Sugawara: 1979:21-3). This finding was further backed up by Phillips-Conroy, et al. in 1991, when they categorized three different types of groups, all of which displayed a certain amount of admixture (Phillips-Conroy, et. al.: 1991:355-6). There were no absolutely "pure" anubis or hamadryas groups either in a morphological or behavioral sense. As far as individuals, according to Phillips-Conroy and Jolly, there is a correlation between individual behavior and morphology. According to their 1986 article, the extent to which hybrids exhibited hamadryas-like behavior was proportional to the "hamadryas-ness" of their appearance (Phillips-Conroy, Jolly: 1986:337). The morphology and behavior of an intergeneric hybrid was described by Moore et al. (1999). The result of a Macaca mulatta x Papio hamadryas cross, this offspring exhibited external features and behaviors of both parents, as well as a vocal sound that appeared to be a combination of both species. Its head, face http://www.soa.ilstu.edu/anthropology/theses/laas/index.htm (3 of 15) [11/30/2007 1:20:42 PM] The Effects of Interbreeding on the Morphological and Genetic Attributes of Non-human Primates / Beth Christine Laas and hair color was more like that of a macaque, while the body build resembled that of a baboon (Moore et al.: 1999:126). Behaviors such as the mantle shake and eyebrow raise were the same as in a baboon, but he lacked the teeth grinding and threat display common to this species. He also displayed fewer vocalizations than both of the parent species (Moore et al. 1999:121). This hybrid is the only long-term survivor of this type of cross, and most of the other infants died within 72 hours (Moore et al.: 1999:121). A cross between a gibbon and a siamang in captivity also produced an intergeneric hybrid. In this case, the offspring was also described as intermediate between each parent. The siabon displayed features that were found in both parents as well as a few that were not found in either. Wolkin and Myers suggested that since the siabon had more features in common with the siamang, but lacked the specialized adaptations (such as the laryngeal sac and alter-developing ischial callosities), that her features are more generalized and more closely resemble an ancestral hylobatid (the siamang is considered more generalized than the gibbon [p. 217]). Hybridization not only occurs between two species in a natural environment. It also has been documented between two different genera: Papio and Macaca. These are often less successful than interspecific crosses, but there is one documented case of a "rheboon" male that survived to reproductive age. "Baboons and macaques are closely related in phylogenetic terms and are quite genetically similar as well" (Moore, et. al.: 1999:119); however, this "rheboon" was the only survivor out of twenty-six attempted crosses and the individual was found to be sterile (Moore, et.