Browsing by Author "Willan, Kenneth Brian Ronald."
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Item The breeding and reproductive biology of the vlei rat Otomys irroratus.(1990) Pillay, Neville.; Willan, Kenneth Brian Ronald.; Meester, Jurgens Anthonie Jansen.The breeding and reproductive biology of the vlei rat Otomys irroratus representing three allopatric populations (Committee's Drift, Hogsback and Karkloof) have been studied in the laboratory. The study attempted to establish whether the three populations differed in respect of selected reproductive parameters, and whether the populations are reproductively isolated from one another. The breeding biology of the Hogsback and Karkloof populations was similar while the Committee's Drift population differed from the other two in respect of its smaller litter size and increased interval between pairing and the production of the first litter. Pre-copulatory behaviour differed among populations, with a gradation of increasing intersexual aggression from Hogsback to Karkloof to Committee's Drift pairs. Postnatal growth and development patterns, as well as male reproductive morphology, were indistinguishable among the populations. Attempts at interpopulation breeding were successful. However, some hybrids died before weaning, while those that survived beyond weaning were sterile, particularly those resulting from cross-matings of Hogsback animals with individuals representing the other populations. Interpopulation pairs displayed higher levels of agonistic interaction than did the pure pairings. Growth and development and the reproductive morphology of male hybrids were indistinguishable from those of the parental populations. Interpopulation reproductive variation in o. irroratus appears to be due to a combination of environmental and phylogenetic constraints. 80th pre- and post-zygotic isolating mechanisms would impede gene flow between the populations should they meet in nature. All populations appear to be undergoing active speciation.Item The evolution and socio-ecology of two populations of the Vlei Rat, Otomys irroratus.(1993) Pillay, Neville.; Cooke, John Anthony.; Willan, Kenneth Brian Ronald.This work investigated two important evolutionary processes - speciation and adaptive variation - in two chromosomally-distinct allopatric Natal Midlands populations of the vlei rat Otomys irroratus. The two populations, at Kamberg and Karkloof, differ in the presence of a tandem fusion between chromosomes seven and 12 in the Kamberg karyotype. Speciation studies considered possible reproductive isolating mechanisms. In studies of adaptive variation, socio-ecological characteristics of both populations were investigated. Data on breeding and postnatal development provided evidence of post-zygotic barriers. Interpopulation pairs had reduced breeding success compared to intrapopulation pairs, and some hybrids died before weaning. Surviving hybrids had reduced growth rates, and almost all were sterile. In tests of pre-mating reproductive isolation, ethological barriers were emphasized. Individuals preferred same-population mates, suggesting the existence of mate recognition, which was achieved by means of population-specific courtship behaviour and communication, particularly olfactory, tactile and visual cues. No evidence of population-specific acoustic signals was found, although acoustic cues were associated with agonistic interaction, complementing other communicatory cues to promote increased aggression during interpopulation pairings. Laboratory studies of behaviour and morphology and field work (trapping and habitat assessment) provided information about socio-ecological parameters. The Kamberg habitat was harsher than the Karkloof one, as revealed by differences in seasonal and spatial availability of food and·cover. Cover was the key determinant of the level of sociability of both populations. Sparse, patchy cover selected for a partially communal social system in Kamberg o. irroratus: females were intrasexually tolerant and males were intrasexually highly aggressive. This, in conjunction with male-biased sexual dimorphism, implied that mating was polygynous. Abundant, uniform cover selected for a dispersed social system in Karkloof o. irroratus: females were intrasexually less tolerant than males. Ritualized aggression between males and a low degree of male-biased sexual dimorphism suggested that male may have overlapping home ranges in nature and that mating is promiscuous. Females possibly mated with dominant males, however. Contrasting social systems suggest that adaptation to local environmental circumstances has occurred in allopatry, and that Kamberg and Karkloof o. irroratus are undergoing adaptive speciation. Post-zygotic and pre-mating reproductive barriers appear to have evolved independently in both populations, and could potentially impede gene flow between the populations should they become sympatric. The presence of the tandem fusion in the Kamberg karyotype which, together with genetically-determined factors, may have caused hybrid sterility, suggests that this population is a chromosomally-determined incipient sibling species.Item Social ecology of Otomys irroratus, Rhabdomys pumilio and Praomys natalensis.(1982) Willan, Kenneth Brian Ronald.; Hickman, G.; Meester, Jurgens Anthonie Jansen.This work sets out to describe the socio-ecological niches of otonya irroratus, Rhabdomys pumilio and Praomys natalensis in the Natal midlands. This objective necessitated a broad-based approach in which aspects of the ecological niches, and the social behaviour and social organization of the three species were investigated in field (habitat and trapping) and laboratory (experimental and observational) studies, during the period January 1976-May 1978. To test the assumption that the commonly sympatric O. irroratus, R. pumilio and P. natalensis are primarily adapted to habitats which differ mainly in respect of water availability, an attempt was made to define their positions on a mesic/xeric continuum. The results suggest that in terms of the parameters measured (responses to water and cover availability, and to food and water deprivation), O. irroratus is more nearly mesically adapted than is R. pumilio, with P. natalensis positioned between the two extremes. However, in the case of P. natalensis it is apparently of overriding importance that this species is adapted to disturbed habitats. The mesic habitats preferred by O. irroratus are often of limited size, but resource availability within these areas is apparently high and stable. Extensive areas of suitable habitat are available to R. pumilio, but availability of resources is seasonally variable. The disturbed habitats preferred by P. natalensis arise unpredictably in nature and, depending on the rate of ecological succession, may be short-lived; hence availability of resources to this species is highly unpredictable. Social behaviour was studied by means of dyadic encounters in the laboratory. Communication in O. irroratus, R. pumilio and P. natalensis appears to be dominated by auditory, visual and olfactory signals respectively, although O. irroratus also has well developed visual signals in its communicatory repertoire. These differences are explained mainly in terms of patterns of diel activity and habitat preferences: R. pumilio is adapted to bright light, and its visual signals are subtle; O. irroratus is adapted to dim light, and its visual signals involve gross changes in posture, while the use of loud, low frequency vocalizations would allow conspecifics to know their precise location in relation to one another in dense vegetation; P. natalensis is nocturnal, and hence appears to emphasize olfactory (and possibly ultrasonic) communication, which would permit transfer of information in the dark. O. irroratus is overtly highly aggressive, but agonism is ritualized; ritualization of aggression has probably evolved to allow high densities of this species to exploit limited areas of prime habitat. R. pumilio is less overtly aggressive than O. irroratus, but aggression tends to be unritualized; direct aggression probably acts as a spacing mechanism, and is possible in view of the large areas of habitat available to this species. Agonism is poorly developed and ritualized in P. natalensis, permitting high sociability and group formation in this species, and maximum exploitation of locally abundant resources. Social organization was studied in the field (trapping studies) and by means of dyadic encounters in the laboratory. The social system of O. irroratus appears to include temporal territoriality, which would permit animals to live in close spatial association (in small areas of habitat) while seldom actually meeting. Hierarchical ranking occurs in male O. irroratus and R. pumilio, with competition in both most likely being for mating opportunities. Breeding females of these two species are intrasexually territorial, in the case of Q. irroratus thereby providing dispersing young with adequate space for establishment of a home-range in prime habitat, and in R. pumilio protecting the young against conspecific female aggression until dispersal. R. pumilio tends to aggregate in mesic refuge habitats in winter, which apparently reflects the seasonality of resource availability to this species in drier environments. P. natalensis is colonial, an adaptation facilitating maximum exploitation of temporarily abundant resources in transitory disturbed habitats. In response to intense courtship by the male, female R. pumilio appear to undergo reflex ovulation; this strategy would maximize the chances of fertilization occurring during occasional meetings between males and females. Courtship intensity is low in P. natalensis, and presumably because the colonial social system of this species ensures frequent association between potential mates, females appear to ovulate spontaneously. Sexual activity was not observed in O. irroratus. The life-history tactics of the three species are such that o. irroratus and P. natalensis respectively appear to be K- and r-selected, with R. pumilio falling between these two extremes on an r-K continuum.Item Systematic revision of the golden mole genera : Amblysomus, Chlorotalpa and Calcochloris (Insectivora : Chrysochloromorpha ; Chrysochloridae)(1995) Bronner, Gary N.; Meester, Jurgens Anthonie Jansen.; Rautenbach, Ignatius Lourens.; Cooke, John Anthony.; Willan, Kenneth Brian Ronald.Patterns of variation in hyoid morphology, chromosomal properties and craniodental characteristics among ten chrysochlorid species from South Africa were studied to clarify generic relationships among taxa assigned variably to Amblysomus, Chlorotalpa and Calcochloris by previous authors. Intra-specific variation in hyoid morphology was negligible, but inter-specific differences were marked. Similarly, intra-specific karyotypic variation was negligible, except in A. hottentotus, which displayed three cytotypes. These data supported the recognition of Chlorotalpa, Calcochloris and Neamblysomus as taxa distinct from Amblysomus. Only one (presence/absence of M3) of the seven dental traits used by previous authors was consistent enough within species to be taxonomically useful in this work. Dental variability within species appeared to arise from the morphological differences between deciduous and permanent teeth, which may occur together in the same toothrow. Intra-specific craniometric variation in most species involved pronounced sexual size dimorphism, but negligible age-related variation. In the more widespread species, patterns of geographic variation were dominated by divergence in overall size, although subtle differences in cranial shape were also evident. Multivariate analyses confirmed the validity of subspecies in Chlorotalpa sclateri and Calcochloris obtusirostris, and showed that A. hottentotus (as traditionally recognized) includes: four cryptic species; five subspecies (including A. h. iris); and several populations that should be relegated to A. corriae. Inter-specific morphometric variation was dominated by overall size. The species fell into two size groups, and eight phena that differed mainly in skull width, palatal shape, rostrum breadth and claw size. Inter-specific relationships suggested by phenetic analyses of metric and mixed-mode data were, however, incongruent owing to discordance between different data suites. Evolutionary relationships inferred by integrating data suites, using either equal or differential weights, indicated that a strong phylogenetic signal was present in the data. Phylogenetic analyses showed that the differentially weighted treatment was more consilient with character-state distributions. A phylogram based on the differential-weights cladogram was used to derive a revised phylogenetic classification for the Chrysochloridae. Unlike previous treatments, this classification affords Carpitalpa and Neamblysomus generic rank, and assigns C. leucorhina from equatorial Africa to Calcochloris, rather than to Chlorotalpa.