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- Poecilia mexicana (4)
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Cave animals are widely recognised as model organisms to study regressive evolutionary processes like the reduction of eyes. In this paper, we report on the regressive evolution of species discrimination in the cave molly, Poecilia mexicana, which, unlike other cave fishes, still has functional eyes. This allowed us to examine the response to both visual and non-visual cues involved in species discrimination. When surface-dwelling females were given a chance to associate with either a conspecific or a swordtail (Xiphophorus hellerii) female, they strongly preferred the conspecific female both when multiple cues and when solely visual cues were available to the female. No association preference was observed when only non-visual cues were provided. In contrast, cave-dwelling females showed no preference under all testing conditions, suggesting that species recognition mechanisms have been reduced. We discuss the role of species discrimination in relation to habitat differences.
Seasonal, host sex and age-related variations in helminth egg and coccidian oocyst counts were investigated in a naturally infected wild bushbuck (Tragelaphus scriptus) population in Queen Elizabeth National Park, western Uganda from April 2000 to February 2002. The prevalence and mean intensity quantified as the number of eggs and oocysts per gram of faeces were taken as a measure of parasite burdens. Host sex and age-related differences in prevalence values were not found but the overall prevalence of Eimeria sp. was significantly higher during the rainy season, and peak counts were recorded either during or soon after a peak rainfall. A similar trend was observed for Moniezia spp., although the results were marginally not significant. There were also no significant differences in mean intensity values, relative to host sex, age or season.
We examined factors predicting female densities within the common home ranges of related females ("clans") in bushbuck Tragelaphus scriptus Pallas, 1776. In this species, each female forms an individual home range, but the home ranges of matrilineal clan members strongly overlap. We found female densities to increase in areas with high canopy cover. Moreover, individual home range sizes tended to decrease with increasing cover. Food plant availability and intruder pressure by two heterospecific competitors did not significantly affect female densities. Apparently, canopy cover is the major limited resource in this species. A possible explanation is that both adult bushbuck and - even more markedly - fawns hide from predators in dense vegetation, in particular in thicket clumps and coalescences. The study shows an effect of habitat properties (eg sufficient canopy cover) on a within-population level in bushbuck, where female densities differ even among proximate clan areas
A free-ranging bushbuck (Tragelaphus scriptus) population was observed over a period of 3 years, thereby enabling a detailed description of the behavioural repertoire of this widespread but barely investigated solitary African antelope species. Agonistic and submissive behaviour patterns are described, among them several hitherto un- described behaviour patterns - such as "escorting", where territorial males guide intruders to the periphery of their territory - and "push-up position", an extreme form of submissive behaviour. Furthermore, we report on behaviour patterns of males and females during mating as well as on behaviour patterns of parents directed towards their offspring. Again, we describe a hitherto unknown behaviour: the protection of calves by adult males, which may be a socio-positive behaviour directed towards their offspring led by kin selection
Atlantic mollies (Poecilia mexicana) inhabit a variety of surface habitats, but they also occur in a sulfur cave in southern Mexico. We examined male mate choice relative to female body size in the cave population and in the most closely related surface-dwelling population from a nearby river. Males from both populations were either light- or dark-reared and could choose between two differently sized females either on the basis of visual cues in light or on the basis of solely nonvisual cues in darkness. Sexual preferences were estimated from the degree of association. Cave molly males always showed a preference for the larger female, both in light and in darkness. Among the surface males, only light-reared males showed a preference in the visual cues test, but not in darkness. In a control experiment, we demonstrated that male association preferences directly translate into actual mating preferences. Apparently, using visual cues for mate choice is the ancestral state in this system, and using nonvisual cues has evolved as a novel trait in the cave population. We discuss the evolution of nonvisual male mate choice in the context of changed environmental conditions, namely the absence of light, hypoxia, and toxic hydrogen sulfide in the cave
We examined chemical communication in male and female European pond turtles (Emys orbicularis). In simultaneous binary choice tests, a focal animal was given a choice between pheromones from a conspecific and a choice chamber containing untreated water. Females did not show a preference, both when male and when female stimuli were presented. On the contrary, males preferred the odor of a female over untreated water, suggesting that males actively search for females. The strength of preference was positively correlated with the body size difference between the female and the focal male, indicating that males prefer to mate with larger females. Female fecundity is positively correlated with female size in E. orbicularis, which may account for male choosiness. No overall preference for the stimulus animal was observed when males were presented cues from another male. However, the strength of preference was negatively correlated with the difference in body size. Males avoided large males, but oriented towards smaller stimulus males. This reflects that males form dominance hierarchies, where large males aggressively attack smaller ones. Far-range chemical communication probably enables males to minimize the risk of costly aggressive interactions. This is, to our knowledge, the first study on the role of chemical cues for inter-and intrasexual communication in the European pond turtle
The communicatory significance of localised defecation sites in bushbuck (Tragelaphus scriptus)
(2006)
Like several other mammals, bushbuck (Tragelaphus scriptus) deposit faeces in specific localised defecation sites (LDS). A previous study has ruled out a function of LDS in the context of parasite avoidance. In this study, we investigated the communicatory significance of LDS. In a free ranging population, we tested whether LDS serve to demarcate home ranges, and/or if LDS are used for communication in a non-territorial context. In both sexes, LDS increased significantly in number towards the periphery of individual home ranges. However, the distribution pattern of LDS, as revealed by a nearest-neighbour mapping technique (nearest distances between LDS), did not support the idea that LDS serve home range/territory-demarcation because LDS did not follow a pattern of minimal nearest-neighbour distances along the margins of home ranges. We found females to urinate more often in LDS than males. Notably, information transfer was most frequent between sending (urinating or defecating) females and receiving/responding males (urination or defecation at places where a female had previously signalled). Our results suggest that LDS mainly serve for inter- sexual communication in bushbuck
The Mexican tetra Astyanax fasciatus has evolved a variety of more or less color- and eyeless cave populations. Here we examined the evolution of the female preference for large male body size within different populations of this species, either surface- or cave-dwelling. Given the choice between visual cues from a large and a small male, females from the surface form as well as females from an eyed cave form showed a strong preference for large males. When only non-visual cues were presented in darkness, the surface females did not prefer either males. Among the six cave populations studied, females of the eyed cave form and females of one of the five eyeless cave populations showed a preference for large males. Apparently, not all cave populations of Astyanax have evolved non-visual mating preferences. We discuss the role of selection by benefits of non-visual mate choice for the evolution of non-visual mating preferences
Bushbuck (Tragelaphus scriptus) often deposit faeces at specific localised defecation sites (LDS). We tested whether LDS have a function in the context of parasite avoidance. In a population of bushbuck in Queen Elizabeth National Park, Uganda, seven radio-collared individuals were observed. We recorded feeding behaviour inside and outside LDS. Furthermore, pasture contamination with gastro-intestinal tract parasites inside and outside LDS was examined. There were significant differences between the expected and the observed feeding rates inside LDS, but, contrary to our prediction, the bushbuck increased their feeding rate inside LDS. There was no significant difference in the parasite contamination of pastures inside and outside LDS. We discuss the hypothesis that LDS mainly serve a social function in bushbuck communities, whereas parasite avoidance seems to play a minor or no role
We investigated whether female association preferences for males are influenced by black spot disease (BSD), a parasite induced change of the host phenotype. We compared three different species of fish: a gynogenetic hybrid species, Poecilia formosa (amazon molly) and two sexual species (Poecilia latipinna and Poecilia mexicana), which were involved in the natural hybridisation leading to the amazon molly. Contrary to their sexual relatives, asexual amazon mollies significantly avoided images of males infected with black spot disease. We propose that amazon molly females have direct fitness benefits from choosing healthy males. The adaptive significance of the preference for BSD-uninfected males in the asexual amazon molly is yet unclear but may involve avoidance of predation or parasite infection as well as increased sperm availability
Life on the edge: hydrogen sulfide and the fish communities of a Mexican cave and surrounding waters
(2006)
Most eucaryotic organisms classified as living in an extreme habitat are invertebrates. Here we report of a fish living in a Mexican cave (Cueva del Azufre) that is rich in highly toxic H2S. We compared the water chemistry and fish communities of the cave and several nearby surface streams. Our study revealed high concentrations of H2S in the cave and its outflow (El Azufre). The concentrations of H2S reach more than 300 mu M inside the cave, which are acutely toxic for most fishes. In both sulfidic habitats, the diversity of fishes was heavily reduced, and Poecilia mexicana was the dominant species indicating that the presence of H2S has an all-or-none effect, permitting only few species to survive in sulfidic habitats. Compared to habitats without H2S, P. mexicana from the cave and the outflow have a significantly lower body condition. Although there are microhabitats with varying concentrations of H2S within the cave, we could not find a higher fish density in areas with lower concentrations of H2S. We discuss that P. mexicana is one of the few extremophile vertebrates. Our study supports the idea that extreme habitats lead to an impoverished species diversity.
Activity patterns and time budgets of bushbuck (Tragelaphus scriptus) were studied in a free-ranging population in Queen Elizabeth National Park, Uganda from August 2000 to January 2002. We investigated differences in activity patterns in relation to daytime, season, sun radiation, moonlight, age and sex. Bushbuck were found to show peak activities around sunrise and at dawn. No difference in the mean activity rates was found between the dry and wet season. Daytime activity was not predicted by differences in sun radiation, nor was nighttime activity predicted by the presence or absence of moonlight. We found the activity of adult territorial males to be strongly positively correlated with that of females, whereas the activity of young-adult non-territorial males was not significantly correlated with the activity of females. This suggests that young-adult males shift their peak activity to phases when adult territorial males are less active.
Males often face strong mating competition by neighboring males in their social environment. A recent study by Plath et al. (Anim Behav 75:21-29, 2008a) has demonstrated that the visual presence of a male competitor (i.e., an audience male) affects the expression of male mating preferences in a poeciliid fish (Poecilia mexicana) with a weaker expression of mating preferences when an audience male observed the focal male. This may be a tactic to reduce sperm competition, since surrounding males likely share intrinsic preferences for female traits or copy mate choice decisions. Here, we examined the hypothesis that a same-sex audience would affect female mate preferences less than male mating preferences. Our hypothesis was based on the assumptions that (1) competition for mates in a fashion that would be comparable in strength to sperm competition or overt male-male aggression is absent among Poecilia females, and (2) P. mexicana females typically form female-biased shoals, such that almost any female mate choice in nature occurs in front of a female audience. Poecilia females (P. mexicana, surface and cave form, and the closely related gynogenetic Poecilia formosa) were given a choice between a large and a small male, and the tests were repeated while a conspecific, a heterospecific, or no audience female (control) was presented. Females spent more time in the neutral zone and, thus, less time near the males during the second part of a trial when an audience was presented, but-consistent with predictions-females showed only slightly weaker expression of mate preferences during the second part of the tests. This decline was not specific to the treatment involving an audience and was significantly weaker than the effect seen in the male sex.
Mate choice is mediated by a range of sensory cues, and assortative mating based on these cues can drive reproductive isolation among diverging populations. A specific feature of mormyrid fish, the electric organ discharge (EOD), is used for electrolocation and intraspecific communication. We hypothesized that the EOD also facilitates assortative mating and ultimately promotes prezygotic reproductive isolation in African weakly electric fishes. Our behavioural experiments using live males as well as EOD playback demonstrated that female mate recognition is influenced by EOD signals and that females are attracted to EOD characteristics of conspecific males. The dual function of the EOD for both foraging and social communication (including mate recognition leading to assortative mating) underlines the importance of electric signal differentiation for the divergence of African weakly electric fishes. Thus, the EOD provides an intriguing mechanism promoting trophic divergence and reproductive isolation between two closely related Campylomormyrus species occurring in sympatry in the lower Congo rapids.
While several authors suggest that bushbuck (Tragelaphus scriptus Pallas) from tropical areas with an approximately bimodal rainfall pattern breed throughout the year, there is also a report of seasonal breeding in this species. In this study, we provide indirect evidence of seasonality in reproduction by analysing behavioural data (e.g. rates of mixed-sex sightings) in a population of bushbuck inhabiting an equatorial savannah ecosystem in western Uganda. Observation rates of mixed-sex sightings were correlated with rainfall patterns. We suggest that peaks in reproductive behaviour following the wet season may be advantageous if calves are born during the next wet season, when fresh vegetation is available.
Savannah areas affected by human activities such as livestock keeping and agriculture are often characterized by shifts in landscape structuring, with a predominance of few(er) habitat types. This is typically accompanied by pronounced changes in the communities of ungulates. The aim of this study was to find out whether shifts in ungulate communities in Lake Mburo National Park (LMNP) are primarily predicted by an alteration in the composition of the preferred habitat types or if more complex interactions between habitat changes and the prevalence of ungulates occur. Monthly road counts were used to establish the number of eleven ungulate species in LMNP and adjacent unprotected Ankole Ranching Scheme. The common duiker (Sylvicapra grimmia campbelliae Gray, 1843) was found in more abundance in disturbed areas, while showing a significant change in habitat use. Common duiker tended to use the vegetation type otherwise used by the bushbuck (Tragelaphus scriptus dama Neumann, 1902). Our results support the claim that the occurrence of ungulates is not only directly affected by the availability of 'suitable' habitats, but behavioural plasticity and competitive exclusion also need to be considered.
We investigated the response of two populations of the barb Garra barreimiae to different light intensities (0.5-2000 lx) from a light source. Adults of both the surface (epigean) and cave (hypogean) G. barreimiae populations show photophobic behavior. A photophobic response in the cave form was seen only at higher light intensities because the cavefish are eyeless and rely on extra-retinal light receptors to detect light. In contrast, juveniles (surface and cave) showed photophilic behavior, and their preference for the photic zone of the test tank decreased with increasing age. We discuss the potential role played by photophobic behavior for the colonization of caves by previously surface- dwelling fishes.
It has been suggested that all species of spiral-horned antelopes (Tragelaphini) lack territoriality. Furthermore, some authors suggested that bushbuck (Tragelpahus scriptus) males form dominance hierarchies. In this study, we investigated the dominance relationships in two groups of free-ranging bushbuck males in Queen Elizabeth National Park, Uganda. Adult males dominated young-adult bachelors and subadult males, but no distinct dominance relationships were found among adult males. Landau's index of linearity revealed no linear dominance hierarchy in the study populations. Our results support the idea that adult males are territorial, and overt aggression is directed almost exclusively towards bachelors that challenge territory holders.
Grouping patterns within the genus Tragelaphus suggest that species inhabiting open areas tend to live in larger groups, while species preferring dense habitats live solitarily or in small family groups. We asked if similar variation would be concealed in the within-species variation of bushbuck (Tragelaphus scriptus). Recent molecular phylogeographic analyses revealed several locally adapted forms of bushbuck in different ecoregions on the African continent. We compared group sizes of south-eastern bushbuck (Tragelaphus sylvaticus) among six different populations ("ecotypes"). To date, most data on the social organization of bushbuck have been collected from only one population in Queen Elizabeth National Park (QENP) in Uganda. This particular population, however, inhabits an unusual-comparatively open-habitat type, while bushbuck otherwise inhabit dense habitats, leaving doubt whether data collected in QENP are representative of the entire species. We, therefore, compared grouping patterns between ecotypes inhabiting rather open habitats (e.g., dianae and haywoodi) and ecotypes occupying rather dense habitats (e.g., massaicus and ornatus). In bachelor groups and in all-female (spinster) groups, single sightings were the most frequent "group type" in all populations examined. We detected no significant difference among ecotypes in relative frequencies of group size categories in the case of bachelor groups. Spinster group sizes were slightly (albeit significantly) smaller in QENP than in all other areas. Moreover, a comparison of two areas inside (low human pursuit) and outside Lake Mburo National Park (high hunting pressure) in Uganda revealed no significant difference in grouping patterns in response to human pursuit (as reported for impala [Aepyceros melampus] inhabiting the same area). Altogether, our results suggest that group sizes in bushbuck are not dependent on the habitat type they inhabit; neither does human nuisance have an impact on grouping patterns. Hence, an "almost solitary" lifestyle appears to be a characteristic of the entire taxon.