This thesis investigated effects of manipulative experience on tool-using ability of tufted capuchins. Two groups of capuchins were tested on variations of a tool-using task, involving use of an object as a tool to dislodge a reward from a tube. The tasks were modelled after those developed by Visalberghi and Limongelli (1994) and Visalberghi and Trinca (1989). One group of monkeys was provided the opportunity to manipulate task materials without reward; the other group was not. Experiment 1 required subjects to push a rod through the tube. Experience with task materials improved capuchins' efficiency, evidenced by faster completion of trials. In Experiment 2, short pipes could be combined to create a tool of sufficient length. Due to procedural problems, results were inconclusive. Experiment 3 required subjects to manufacture the appropriate tool. Experience with task materials improved capuchins' performance, evidenced by faster completion of trials, less frequent performance of inefficient behaviours, and decrease in errors across trials. When capuchins' performance was compared with enculturated chimpanzees tested previously on the task (Visalberghi, Fragaszy, and Savage-Rumbaugh, 1995), experienced capuchins performed as efficiently after 15 trials as had chimpanzees originally. In Experiment 4, subjects had to dislodge the reward from a tube containing a trap. Because successful levels of performance were not reached, results were inconclusive. In Experiments 5 and 6, appreciation of object affordances was examined. The capuchins demonstrated an ability to distinguish between functional characteristics of objects. These results suggest previous claims regarding limitations of capuchin tool-use may have underestimated their abilities. Further, because object experience enhances tool-using ability, previous comparisons of capuchins with enculturated chimpanzees seem to have misrepresented the magnitude of difference in their abilities. However, future research comparing species with the same experiential backgrounds is necessary to elaborate on differences in cognitive processes underlying capuchin and chimpanzee tool-using behaviour.

It is hypothesised that young baboons are potentially vulnerable to the effects of seasonal stress. Data were collected on the behaviour of this age group during a 12 month field study of a troop of olive baboons (Papio anubis) on the Laikipia plateau, Kenya, using a hand-held computer and a new program written by the author. Long-term environmental records for this site were continued. At this site there is a seasonal pattern of rainfall with inter-annual variation. Measures of biomass indicate that there are seasonal fluctuations in baboon food availability. Patterns in the occurrence of one component of the baboon's diet, Acacia species, are presented. The varying behaviour of the troop as a whole is related to food availability. Differential use of the home range and observed sub-trooping behaviour are interpreted as adaptive strategies for living in a seasonal environment. Developmental change in the behaviour associated with the mother-offspring relationship is described. Patterns in the time spent in contact with and close proximity to the mother from this site are compared with those from other sites, and striking similarities are found. Many of the behavioural activities of infants and young juveniles, i.e. feeding, moving, types of exploring, visually attending, grooming, and receiving affiliative approaches, exhibit patterns of developmental change. Interactions between activities are examined in the context of the complete activity budget. The effect of seasonal stress on young baboons is examined by using a technique of curve fitting. Data are compared between periods of higher and lower food availability. Significantly more time is spent feeding and less time spent in social and attending activities in the 'worst' than the 'best' months. Young baboons vary their diets seasonally. There is evidence that older infants (weanlings) are more vulnerable to the impact of seasonal stress than either young infants or young juveniles.

This thesis investigates the nature of post-conflict behaviour and relationships among individuals in two captive groups of pigtail macaques, Macaca nemestrina, and one wild troop of olive baboons, Papio anubis. Data were collected during periods of seven and twelve months respectively. In both pigtail groups, conflicts were more frequently reconciled between opponents with strong affiliative ties. However, reconciliation was twice as common in the well-established group where individuals' social networks were more compact. It is argued that the more intense ties produced by restricted networks increased the likelihood of reconciliation. Reconciliation was demonstrated among wild olive baboons, occurring at a rate consistent with a relatively intolerant dominance style. Opponents who were close kin or of similar rank reconciled relatively frequently, but reconciliation rarely followed conflicts associated with food. Olive baboons did not 'console' each other, consistent with the hypothesis that consolation requires an ability to empathise with victim distress. Initiation of post-conflict attacks on third parties was not elevated in victims of aggression. Among the baboons, both victims and initiators of aggression exhibited elevated rates of post-conflict self-directed behaviour (a combined measure of scratching, autogrooming, body-shaking and yawning). Reconciliation reduced both SDB and the incidence of further aggression. However, reconciliation only reduced SDB among individuals involved in conflicts in which they had both received and delivered aggression. Female baboons showed significantly higher rates of SDB when their nearest neighbour (within 5 m) was a dominant conspecific than when he or she was a subordinate individual, supporting the hypothesis that SDB indexes stress in primates. This result suggests that SDB can be used to index relationship security in primates.

The problems of what primates communicate with their vocalizations and to what extent they refer to events in the external environment are raised in numerous studies of primate vocal communication. To investigate these issues, I concentrate on the calls of two primate species with a similar fission-fusion social organization. I report here the results of a one year field study on the Central American spider monkey (Ateles geoffroyi frontatus) and of a captive study on the chimpanzee (Pan troglodytes). I collected observational data and conducted playback experiments on two types of calls of the spider monkey, whinnies and alarm barks. Spider monkeys use two functionally distinct whinnies, a feeding whinny and a locational whinny. Detailed acoustic analyses of whinnies given in different contexts showed that in three females an acoustic feature-number of arches in the fundamental frequency of the call- could be used to differentiate between them. Although the whinnies of different monkeys within the same community exhibit enough acoustical differences for individual vocal recognition to take place, spider monkeys did not discriminate familiar individuals' whinnies from those of strangers from another community. In predator contexts spider monkeys use barks to attract other conspecifics to a site. However, my data do not support the existence of two types of alarm barks for aerial versus terrestrial predators. I conducted two different types of experiments on the calling behaviour of captive chimpanzees in response to food-finding. I examined whether the quantity and divisibility of food, or the presence of an audience, influence calling behaviour. Chimpanzees' food- associated calls, i.e. rough grunts, functioned to indicate food availability, and they were produced or suppressed depending on how shareable the food was and whether or not other individuals were nearly. Several spider monkeys' and chimpanzees' calls have the ability to function referentially, but cannot be considered devoid of a motivational content.

Die Evolution der heutigen globalen Diversität wurde in den letzten Millionen Jahren insbesondere durch klimatische Schwankungen und entsprechende Veränderungen biologischer Lebensräume beeinflusst. Besonders deutlich ist der Einfluss von Glazialen und Interglazialen auf die Evolution von Organismen der Nördlichen Hemisphäre. Weniger klar hingegen ist, inwiefern sich diese klimatischen Verhältnisse auf (sub-) tropische Regionen ausgewirkt haben, insbesondere auf das Afrikanische Savannen Biom. Unabhängig davon, haben Umweltveränderungen in tropischen und nicht-tropischen Gebieten zur Entstehung vieler der gegenwärtigen Hybridzonen geführt, in denen zuvor geographisch separierte Populationen in sekundären Kontakt gekommen sind. Obwohl Hybridisierung im Tierreich inzwischen nicht mehr als ein rares Phänomen betrachtet wird, ist das tatsächliche Ausmaß und die Bedeutung von Hybridisierung in der Evolution von Tieren noch lange nicht vollständig geklärt. Die Verbreitung Grüner Meerkatzen der Gattung Chlorocebus reflektiert nahezu die Ausdehnung Afrikanischer Savannengebiete und basierend auf Beobachtungen im Freiland sowie auf morphologischen Merkmalen von Museumsexemplaren, hybridisieren die meisten der sechs anerkannten parapatrischen Arten in ihren jeweiligen Kontaktzonen. Aufgrund dieser Eigenschaften habe ich in meiner Doktorarbeit Grüne Meerkatzen als Modellsystem genutzt, um zum einen die Bedeutung von Hybridisierung in der Evolution von Primaten und Tieren im Allgemeinen zu untersuchen, und zum anderen, um wesentliche Trends in der Evolution von Savannensäugetieren zu analysieren. Um in einem ersten Schritt grundlegende Informationen über die mitochondriale Diversität und Verbreitung der Grünen Meerkatzen zu erlangen, generierte ich vollständige Cytochrom b Sequenzen von Proben, die alle sechs Arten der Gattung und große Teile der gesamten Verbreitung Grüner Meerkatzen repräsentierten. Des Weiteren nutzte ich Sequenz-Informationen zweier Y-chromosomaler Loci, ein Fragment der sex determining region (SRY) und das letzte Intron des Zinc finger (ZFY), um eventuell zeitlich zurückliegende Hybridisierungsereignisse nachzuweisen. Um räumliche sowie zeitliche phylogeographische Muster zu rekonstruieren, habe ich basierend auf den bisher gewonnen Daten weitere mitochondriale Marker von selektiven Proben aller mitochondrialer Kladen sequenziert. Abschließend habe ich die Phylogeographie Grüner Meerkatzen mit den Phylogeographien dreier anderer weit verbreiteter Savannensäugetier-Gattungen verglichen, mit Pavianen (Papio), Warzenschweinen (Phacochoerus), und Kuhantilopen (Alcelaphus). Meine Analysen der mitochondrialen Daten lassen neun klar abgegrenzte Kladen erkennen, die keiner bisher vorgeschlagenen Taxonomie entsprechen. Zahlreiche para- und polyphyletische Beziehungen, verursacht durch nicht übereinstimmende Verbreitungsmuster mitochondrialer Kladen und morphologischer Merkmale, liefern Hinweise auf anhaltende introgressive Hybridisierung in den Kontaktzonen aller Arten, mit Ausnahme der beiden westafrikanischen Arten. Darüber hinaus weisen die Ergebnisse der mitochondrialen Analysen auf potentiell vergangene introgressive Hybridisierungsereignisse hin, die geographisch nicht in Gebiete gegenwärtiger Kontaktzonen fallen. Dies kann im Falle von C. pygerythrus in Ostafrika anhand der Y-chromosomalen Daten bestätigt werden. Männchen basierte Introgression und nuclear swamping haben hier offensichtlich zu dem zytonukleären Aussterben eines historischen Taxons geführt. Aber nicht alle diskordante Muster in der mitochondrialen Phylogenie sind Anzeichen für zurückliegende introgressive Hybridisierung. Innerhalb der Verbreitung von C. tantalus weisen sowohl mitochondriale als auch Y-chromsomale Daten auf zwei klar separierte und morphologisch kryptische Taxa hin (ein westliches und ein östliches Taxon), ein Befund der möglicherweise auf das Vorhandensein einer neuen Grünen Meerkatzenart hindeutet. In Übereinstimmung mit mitochondrialen Ergebnissen weisen Y-chromosomale Daten ebenfalls keine Anzeichen für Hybridisierung zwischen C. sabaeus und der westlichen C. tantalus-Form in Westafrika auf. Entgegen früherer Annahmen stellt der Volta Fluss und seine nördlicheren Zuflüsse offenbar auf gesamter Länge eine geographische Barriere dar und nicht wie vorher angenommen nur im südlicheren Teil des Flussverlaufs. Die Phylogeographie Grüner Meerkatzen weist somit auf eine komplexe evolutionäre Geschichte hin. Aufgrund der phylogenetischen Rekonstruktionen und Datierungen gehe ich von einem Westafrikanischen Ursprung der Gattung vor ca. 2,46 Millionen Jahren aus. Des Weiteren liefert die Phylogenie Hinweise auf eine erst später folgende Ausbreitung bis nach Südafrika und auf zwei zeitlich getrennte Besiedelungen nordöstlicher Regionen, eine vom Westen und eine von eher südlicheren Regionen aus. Im Vergleich zu den anderen Savannensäugetieren können keine zeitlichen Übereinstimmungen in Aufspaltungsmustern gefunden werden, weder im Vergleich zwischen den Primaten noch zwischen den Ungulaten. Zudem fallen Aufspaltungen innerhalb der Gattungen zeitlich sowohl mit kalt-ariden als auch mit warm-humiden Perioden zusammen. Veränderungen in Populationsgrößen innerhalb der letzten 500.000 Jahre zeigen unterschiedliche Muster zwischen Primaten und Ungulaten, die wahrscheinlich mit verschiedenen ökologischen Anpassungen und Habitat Präferenzen zu erklären sind. Da außerdem keine klaren Zusammenhänge zwischen dem zeitlichen Auftreten von Populationsschwankungen und dem letzten Interglazialem bzw. Glazialen Maximum gefunden werden können, scheint ein Einfluss von regional geprägtem Klima in Afrika wahrscheinlicher. Zusammengefasst hat die Evolution Grüner Meerkatzen im frühen Pleistozän begonnen und klimatische Schwankungen im Quartär haben vermutlich zu wiederkehrenden Veränderungen in der Ausbreitung geführt. Diese begünstigte die Entstehung von sekundären Kontaktzonen und führte zu weit verbreiteter introgressiver Hybridisierung innerhalb der Gattung. Das zytonukleäre Aussterben ehemaliger Taxa oder Populationen als Ergebnis lang anhaltender introgressiver Hybridisierung und darauffolgendem nuclear swamping machen zudem den potentiellen Einfluss von Hybridisierung in der Evolution von Primaten und Tieren im Allgemeinen deutlich. Die Phylogeographien der Savannensäugetiere zeigen, dass Aufspaltungen innerhalb der Gattungen sehr wahrscheinlich durch ausgeprägt humide wie auch aride Bedingungen begünstigt wurden, die eher durch regionale Klimatische Veränderungen in Afrika zu erklären sind als durch Glaziale Zyklen und Klimaschwankungen der Nördlichen Hemisphäre. Die Ergebnisse meiner Doktorarbeit betonen die Notwendigkeit der Analyse von sowohl mütterlich als auch väterlich vererbten Markern in phylogeographischen Studien, um ein möglichst vollständiges Bild von evolutionären Prozessen, Hybridisierung, sowie von genetischer und taxonomischer Vielfalt zu erlangen.

Summary: This dissertation is focused on observation of locomotion two groups of mantled guereza in ZOO Prague and ZOO Usti nad Labem. The observation took place in the summer of 2011. The locomotion behavior is for better clarity presented by tables, diagrams and detailed description of the breeding facilities of both groups. The work contains list of all species of guereza, including all subspecies of mantled guereza. Key words: primates, old world monkeys, Colobinae, black and white colobus monkey, mantled guereza, threat, behavior, breeding in the zoo.

abstract: Dietary diversity is an important component of species’s ecology that often relates to species’s abundance and geographic distribution. Additionally, dietary diversity is involved in many hypotheses regarding the geographic distribution and evolutionary fate of fossil primates. However, in taxa such as primates with relatively generalized morphology and diets, a method for approximating dietary diversity in fossil species is lacking. One method that has shown promise in approximating dietary diversity is dental microwear analyses. Dental microwear variance has been used to infer dietary variation in fossil species, but a strong link between variation in microwear and variation in diet is lacking. This dissertation presents data testing the hypotheses that species with greater variation in dental microwear textures have greater annual, seasonal, or monthly dietary diversity. Dental microwear texture scans were collected from Phase II facets of first and second molars from 309 museum specimens of eight species of extant African Old World monkeys (Cercopithecidae; n = 9 to 74) with differing dietary diversity. Dietary diversity was calculated based on food category consumption frequency at study sites of wild populations. Variation in the individual microwear variables complexity (Asfc) and scale of maximum complexity (Smc) distinguished groups that were consistent with differences in annual dietary diversity, but other variables did not distinguish such groups. The overall variance in microwear variables for each species in this sample was also significantly correlated with the species’s annual dietary diversity. However, the overall variance in microwear variables was more strongly correlated with annual frequencies of fruit and foliage consumption. Although some variation due to seasonal and geographic differences among individuals was present, this variation was small in comparison to the variation among species. Finally, no association was found between short-term monthly dietary variation and variation in microwear textures. These results suggest that greater variation in microwear textures is correlated with greater annual dietary diversity in Cercopithecidae, but that variation may be more closely related to the frequencies of fruit and foliage in the diet. / Dissertation/Thesis / Doctoral Dissertation Anthropology 2015

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The aim of the present work was to investigate the impact of the varying environmental conditions on the taxonomic and phenotypic diversification of a geographically widespread and ecologically successful Old World primate genus, the macaques (Cercopithecidae: Macaca). To this end, the relationship between geography, ecology, phylogeny, and phenotypic variation among macaques was investigated. Constraints to phenotypic variation – and thus evolution – were also analysed in the form of observed amounts of phenotypic variation and patterns of phenotypic integration. A total of 72 standard linear measurements of teeth and associated cranial and mandibular structures were taken for a total sample of 744 specimens from 13 species of macaques. Climate and ecological data were collated from the literature. Univariate and multivariate statistics were employed for the analysis. Patterns of variation, covariation, and allometry were analysed in the dentition, both within and between species. The ecogeographical analysis was carried out by means of two-block partial least squares and a type of multivariate regression, both in a phylogenetic framework. Phylogenetic signal was tested for by means of Blomberg’s K. Macaque teeth differ in their variability. All teeth covary with each other, although correlations are strongest within tooth classes. Size was a strong contributing factor to dental integration, as evinced by lower correlations between teeth once allometric effects were removed. Integration patterns also showed modularity between the anterior and the posterior dentition. Between-species variation in overall craniodental size was associated with temperature, latitude, and body size. Species also varied, albeit to a lesser degree, along an antero-posterior contrast in relative tooth size. Larger anterior were found to be associated with frugivory and tropical ecology, whereas a larger posterior dentition was linked to a more folivorous diet and temperate environments. The latter pattern was largely a function of phylogenetic relatedness. Phylogenetic signal was generally strong in the dentition, although it was substantially greater in the anterior teeth (incisors and canines) than in the posterior teeth (premolars and molars). Macaques show adaptive differentiation in body size in response to temperature along a latitudinal cline, corroborating the presence of the Bergmann effect in macaques. There was no conclusive support for further adaptive differentiation, despite an association between relative tooth size and diet. Allometry appears to channel evolutionary divergence of macaques along a line of least evolutionary resistance, and developmental modularity allows for partly uncoupled evolution of the anterior and posterior dentition. Future research should be aimed at broadening the taxonomic scope to include craniodental variation of the African papionins and cercopithecins in order to put the observed macaque patterns in a broader evolutionary context.