Habitat segregation among mimetic ithomiine butterflies (Nymphalidae)

The extensive wing pattern diversity observed among sympatric unpalatable mimetic butterflies is difficult to explain. Diversity is a paradox because selection by predators is expected to drive local species to use the same aposematic patterns. Habitat segregation among mimicry complexes has been suggested as a hypothesis to explain how diversity could be maintained. However, very few studies have tested this hypothesis. To test whether mimicry complexes are associated with particular habitats, I sampled a diverse assemblage of ithomiine butterflies from eastern Ecuador comprising nine discrete mimicry complexes. Butterflies were sampled in four habitats varying along a gradient of succession. A total of 43 species and 902 individuals were sampled. Ithomiine species richness and abundance were lowest in open habitats, and habitat preferences were documented for many species. Mimicry complexes exhibited significant habitat differences supporting the role of habitat segregation in maintaining mimetic diversity. However, there was obvious overlap among mimicry complexes, particularly involving the two numerically dominant patterns at the site. The pattern of segregation appears to be driven by common species, with relatively little evidence that the distribution of rarer species matches that of the more abundant species. Thus, habitat segregation is likely to play a role in the evolution of mimetic diversity as a result of segregated abundant model species, but the effect is probably weak and other factors are also important.     

Fitness and habitat segregation of British tree-hole mosquitoes

Abstract.

• 1 Populations of three mosquitoes, Aedes geniculutus (Olivier), Anopheles plumbeus Stephens and Culex torrentium Martini, were found in three types of tree holes: rot holes penetrating the bark of the host tree and exposed, or deeper, more cryptic, bark-lined buttress holes. Aedes geniculatus occupied all three tree-hole types but reached their greatest abundance in exposed buttress holes; A.plumbeus predominated in rot holes; C.torrentium occupied buttress holes exclusively.
• 2 After a 1 month period of winter freezing, larvae of A.plumbeus survived better than larvae of A. genicularus. Freeze resistance of A.plumbeus increased from cryptic buttress to rot holes. Freeze resistance of A.geniculatus declined from exposed buttress to cryptic buttress to rot holes and was markedly higher in more protected than in more exposed macrohabitats.
• 3 Among manipulated larval cohorts of A.geniculutus and A.plumbeus, survivorship, pupation success, pupal weight, and/or biomass yield did not differ among tree-hole types or was the reverse of freeze-related survivorship. Despite seasonal differences in fitness correlates, annual fitness may be similar among tree-hole types for both species. Manipulations carried out at a single census or during a single season are therefore likely to produce misleading information about fitness variation among sub-habitats.
• 4 Both species should exploit a broad range of tree-hole types but A.plumbeus occupies more restricted types of tree holes than does A.geniculutus. The restricted habitat usage by A.plumbeus does not parallel their fitness among tree-hole types in southern Britain but does resemble that of other tree-hole anophelenes. We therefore propose that habitat selection by A.plumbeus is more likely due to their phylogenetically determined physiological tolerances and behaviour than to competition from coexisting A.geniculatus.

     

Habitat selection and aggression as determinants of spatial segregation among damselfish on a coral reef

Adults of many closely related coral reef fish species are segregated along gradients of depth or habitat structure. Both habitat selection by new settlers and subsequent competitive interactions can potentially produce such patterns, but their relative importance is unclear. This study examines the potential roles of habitat selection and aggression in determining the spatial distribution of adults and juveniles of four highly aggressive damselfishes at Lizard Island, northern Great Barrier Reef. Dischistodus perspicillatus, D. prosopotaenia, D. melanotus, and D. pseudochrysopoecilus maintain almost non-overlapping distributions across reef zones, with adults of one species dominating each reef zone. Juveniles exhibit slightly broader distributional patterns suggesting that subsequent interactions reduce overlap among species. Although habitat choice experiments in aquaria suggest that associations between juveniles and substrata types in the field are partly due to habitat selection, large overlaps in the use of substrata by the different species were also found, suggesting that substratum selection alone is insufficient in explaining the discrete spatial distributions of adults. The strength of aggressive interactions among all four species was tested by a "bottle" experiment, in which an adult or juvenile of each species was placed in the territories of adult fish on the reef. The greatest levels of interspecific aggression were directed against adults and juveniles of neighbouring species. The highest levels of aggression were associated with species exhibiting the greatest levels of overlap in resource use. Evidently both habitat selection and interspecific aggression combine to determine the adult distributions of these species.     

Habitat segregation by fishes in western Taiwan rivers

Habitat segregation has been recognized as an important means of resource partitioning in river fish assemblages. Association between river fishes and habitat features were identified through principal component analysis (PCA). Fish and habitat data were collected in the Kaoping, Tsengwen, Choshui, and Tatu rivers from July 1997 to June 1999 in western Taiwan. Electrofishing was used to collect fishes in grids, and environment variables in the sampled areas were measured immediately after sampling. The first four PCs of depth, velocity, cover, substrate and water temperature variables explained 92.3% of the total variance in habitat characteristics. Varicorhinus barbatulus, Acrossocheilus formosanus and Tilapia mossambica in the upper right quadrant, indicated preference for deep‐water habitat. Species usually associated with shallow water habitat include several minnow species. Rhinogobius brunneus, Hemimyzon formosanum, Sinogastromyzon puliensis and Crossostoma lacustre distribution is centred in the upper and lower left, indicating preference for shallow water. Principal component (PC) scores on Zacco pachycephalus, Abbottina brevirostris and Hemiculter leucisculus appeared to have no special preference. Differences in habitat use exhibited by these species may be related to success in finding food and avoiding predators.     

Linkage relationships among four enzyme loci in the predatory, treehole mosquito

Estimates are reported for the linkage distances between the isozyme loci for hydroxybutyrate dehydrogenase (HbDH), phosphoglucomutase (PGM), phosphoglucose isomerase (PGI), and esterase 8 (Est 8) in the predatory, treehole mosquito, Toxorhynchites rutilus rutilus. Measured linkage distances suggest that these four loci form a single autosomal linkage group approximately 47.4 linkage units in length. The estimates were measured with error because of small family sizes and, perhaps, because of different rates of recombination between males and females or the presence of a paracentric inversion associated with this linkage group. The four loci are tentatively placed in the following order: Est 8--HbDH--PGM--PGI.     

Plant coexistence in coastal heaths: Habitat segregation in the post-fire environment

Abstract Previous studies of heaths on Pleistocene coastal sands showed consistent variation in patterns of floristic composition between areas on ridges and slopes but not within them. A large wild-fire that swept this system provided an opportunity to observe temporal processes in habitat segregation of species. Ridges and slopes were found to differ in species richness of seedlings in 0.0625 m2 quadrats and there was no evidence that the species richness in the two habitats was converging over 3 years. This suggests that initially these processes differ in space between the two habitats with more species in the wet heath being packed into a smaller area than in the dry heath. Seed-banks of species did not saturate available space for recruitment in either habitat, but seedling densities differed asymmetrically between habitats across two pairs of species studied. Experimental manipulation of seeds among habitats also showed distinct differences in establishment and survival among representative species from different habitats. The presence of seed-banks of wet-heath species in dry heath suggested that recruitment of their seedlings may occur there, but experiments showed that seedlings that arise from them do not survive there. Conversely, although no seed-bank of either dry-heath species studied was found in wet heath, our manipulations showed that their seedlings could grow there. Thus, superficially, physiological tolerance appeared to limit regeneration of wet-heath species in dry heath. However, there appeared to be no physiological limit for dry-heath plants to survive in wet heath. This paper shows that segregation of species may operate at either of two stages of the life cycle: at dispersal when safe sites for establishment are required, and at recruitment after seedlings have established. These findings highlight the importance of the regeneration niche in structuring community composition in coastal heaths, and contrast with traditional explanations of species segregation invoking physiological tolerance and competition at later life-history stages.     

Habitat segregation and interactive effects of multiple predators on a prey assemblage

1. In a field experiment we examined the interactive effects of two common predators of zooplankton, bluegill sunfish (Lepomis macrochirus) and Chaoborus spp. on the growth rate and habitat use of three congeneric prey species (Daphnia). Bluegill and Chaoborus both consume Daphnia, but bluegill also prey on Chaoborus. The prey species, Daphnia pulicaria, D. rosea and D. retrocurva, differed in body size and vertical distribution. We expected the largest species, D. pulicaria, to be most vulnerable to fish predation and the smallest species, D. retrocurva, to be most vulnerable to Chaoborus predation. 2. As we expected, the population growth rate of D. pulicaria was significantly reduced by fish. However, Chaoborus also significantly reduced the growth rate of this species. No significant interaction effect was detected, indicating that the effect of these predators was additive. The growth rates of D. rosea and D. retrocurva were significantly reduced by Chaoborus, but a significant interaction effect indicated that the effect of Chaoborus was stronger in the absence of fish than when fish were present. Therefore the impact of Chaoborus and fish on D. rosea and D. retrocurva was non-additive. The interactive effect of the two predators on D. retrocurva was greater in magnitude than on D. rosea. 3. In the absence of predators, the three Daphnia species showed no differences in mean habitat depth between day and night. Both predators significantly affected diel habitat use of D. pulicaria and D. rosea. Fish caused both of these Daphnia species to move deeper during the day, whereas Chaoborus caused Daphnia to move into shallower water at night. Daphnia retrocurva tended to migrate upwards at night in all predator treatments, but no significant differences in migration were observed among the predator treatments. The effects of predators on habitat use were not interactive for any prey species. 4. Our results suggest that body size, habitat use and the diel migratory response to predators are important factors mediating the interactive effects of multiple predator types on zooplankton.     

Habitat overlap/niche segregation in two Umbilicaria lichens: a possible mechanism

Summary Umbilicaria muhlenbergii and Umbilicaria papulosa are two similar lichens which show extensive habitat overlap. Current theory predicts that for those resources that are limited, and of critical importance to both species, segregation is demand for or use of those resources ought to be present. Controlled experiments showed that the upper and lower cortical surfaces of the two species functioned in very different ways so that U. muhlenbergii is wetted most rapidly by runoff and mist-like precipitation whereas U. papulosa is wetted most rapidly by large raindrops. An analysis of the meteorological conditions in the natural habitat for the two lichens shows that the four months providing significant productivity in U. papulosa are the same months with a very high frequency of thunderstorms (which produce large drops). Conversely, the months which produce the greatest production in U. muhlenbergii are spread out over most of the year. When water is supplied after periods of dryness, the two species absorb it at different times and from different sources. This difference supports current ideas about niche segregation in plants but represents the first time it has been demonstrated for lichens.     

Diversity among African pygmies

Although dissimilarities in cranial and post-cranial morphology among African pygmies groups have been recognized, comparative studies on skull morphology usually pull all pygmies together assuming that morphological characters are similar among them and different with respect to other populations. The main aim of this study is to compare cranial morphology between African pygmies and non-pygmies populations from Equatorial Africa derived from both the Eastern and the Western regions in order to test if the greatest morphological difference is obtained in the comparison between pygmies and non-pygmies. Thirty three-dimensional (3D) landmarks registered with Microscribe in four cranial samples (Western and Eastern pygmies and non-pygmies) were obtained. Multivariate analysis (generalized Procrustes analysis, Mahalanobis distances, multivariate regression) and complementary dimensions of size were evaluated with ANOVA and post hoc LSD. Results suggest that important cranial shape differentiation does occur between pygmies and non-pygmies but also between Eastern and Western populations and that size changes and allometries do not affect similarly Eastern and Western pygmies. Therefore, our findings raise serious doubt about the fact to consider African pygmies as a homogenous group in studies on skull morphology. Differences in cranial morphology among pygmies would suggest differentiation after divergence. Although not directly related to skull differentiation, the diversity among pygmies would probably suggest that the process responsible for reduced stature occurred after the split of the ancestors of modern Eastern and Western pygmies.     

Niche segregation among sympatric Amazonian teiid lizards

We examined standard niche axes (time, place, and food) for three sympatric teiid lizards in the Amazon rain forest. Activity times during the day were similar among species. Ameiva ameiva were in more open microhabitats and had higher body temperatures compared with the two species of Kentropyx. Microhabitat overlaps were low and not significantly different from simulations based on Monte Carlo analysis. Grasshoppers, crickets, and spiders were important in the diets of all three species and many relatively abundant prey were infrequently eaten (e.g., ants). Dietary overlaps were most similar between the two species of Kentropyx even though microhabitat overlaps were relatively low. A Monte Carlo analysis on prey types revealed that dietary overlaps were higher at all ranks than simulated overlaps indicating that use of prey is not random. Although prey size was correlated with lizard body size, there were no species differences in adjusted prey size. A. ameiva ate more prey items at a given body size than either species of Kentropyx. Body size varies among species, with A. ameiva being the largest and K. altamazonica the smallest. The two species of Kentropyx are most distant morphologically, with A. ameiva intermediate. The most distant species morphologically are the most similar in terms of prey types. A morphological analysis including 15 species from four genera revealed patterns of covariation that reflected phylogenetic affinities (i.e., taxonomic patterns are evident). A cluster analysis revealed that A. ameiva, K. pelviceps, and K. altamazonica were in the same morphological group and that within that group, A. ameiva differed from the rest of the species. In addition, K. pelviceps and K. altamazonica were distinguishable from other species of Kentropyx based on morphology. Received: 26 December 1998 / Accepted: 15 September 1999     

Distribution of African malaria mosquitoes belonging to the Anopheles gambiae complex

The distribution of malaria vector mosquitoes, especially those belonging to species complexes that contain non-vector species, is important for strategic planning of malaria control programmes. Geographical information systems have allowed researchers to visualize distribution data on maps together with environmental parameters, such as rainfall and temperature. Here, Maureen Coetzee, Marlies Craig and David le Sueur review our current knowledge on the distribution of the members of the Anopheles gambiae complex.     

Trade-offs of predation and foraging explain sexual segregation in African buffalo

1. Many studies have investigated why males and females segregate spatially in sexually dimorphic species. These studies have focused primarily on temperate zone ungulates in areas lacking intact predator communities, and few have directly assessed predation rates in different social environments. 2. Data on the movement, social affiliation, mortality and foraging of radio-collared African buffalo (Syncerus caffer) were collected from 2001-06 in the Kruger National Park, South Africa. 3. The vast majority of mortality events were due to lion (Panthera leo) predation, and the mortality hazard associated with being an adult male buffalo in a male-only 'bachelor' group was almost four times higher than for adult females in mixed herds. The mortality rates of adult males and females within mixed herds were not statistically different. Mortality sites of male and female buffalo were in areas of low visibility similar to those used by bachelor groups, while mixed herds tended to use more open habitats. 4. Males in bachelor groups ate similar or higher quality food (as indexed by percentage faecal nitrogen), and moved almost a third less distance per day compared with mixed herds. As a result, males in bachelor groups gained more body condition than did males in breeding herds. 5. Recent comparative analyses suggest the activity-budget hypothesis as a common underlying cause of social segregation. However, our intensive study, in an area with an intact predator community showed that male and female buffalo segregated by habitat and supported the predation-risk hypothesis. Male African buffalo appear to trade increased predation risk for additional energy gains in bachelor groups, which presumably leads to increased reproductive success.     

Habitat segregation in four species of adult tiger beetles (Coleoptera: Cicindelidae)

ABSTRACT.

• 1 Habitat segregation in four species of tiger beetles, Cicindela cancellata Dejean, C.cardoni Fleutians, C.minuta Olivier and C.sumatrensis Herbst in a river bank ecosystem was studied in dry and wet seasons.
• 2 The four species segregated distinctly along the river beds into separate habitats, with occasional overlapping in both the seasons.
• 3 Among the habitat characteristics considered, vegetation, soil moisture and available prey-size were found to be important in species segregation.
• 4 D ²-analysis showed that the habitat preferences of C.cardoni and C.cancellata were closer to that of C.sumatrensis. The habitat of C.minuta was distinctly separated.
• 5 There was a positive correlation between the mandible length of each tiger beetle species and the length of prey captured.
• 6 Variance in mandible length within species was related to prey-length distribution pattern and to variance in habitats.

     

Resource limitation,habitat segregation,and species interactions of british tree-hole mosquitoes in nature

Summary The insect fauna of water-filled tree holes in southern Britain consists primarily of the mosquitoes Aedes geniculatus, Anopheles plumbeus, Culex torrentium, and a benthic detritivorous fauna that includes primarily the scirtid beetle Prionocyphon serricornis and the chironomid midge Metriocnemus martinii. Culex torrentium has been documented only relatively recently in tree holes but all three species of mosquitoes partition the resource in space and time. When mosquito larvae were forced to coexist in natural tree holes at limiting densities and at higher than natural levels of interspecific encounter, there was no evidence that Aedes geniculatus or Anopheles plumbeus affected pupation success, pupal weight, or development time of the other or that either Aedes geniculatus or C. torrentium affected the survivorship, pupation success, pupal weight, and biomass yield of the other. When A. geniculatus at limiting densities were forced in natural tree holes to live without or to coexist with natural or twice natural densities of P. serricornis and M. martinii, the presence, absence, or superabundance of the benthic insects did not affect pupation success or pupal weight of A. geniculatus; development time of A. geniculatus was faster when a superabundance of the benthic fauna was present. Effects of the benthic fauna on A. geniculatus are slight and the only significant interaction is facilitative, not competitive. The pattern of habitat segregation among treehole mosquitoes in southern Britain is characteristic of their respective genera and we propose that this pattern is more likely (but not certain) to have arisen through a process of independent evolution than through competitively driven niche shifts among already coexisting species.     

Genetic structure among Thai populations of Aedes aegypti mosquitoes

Thirty‐one field populations of Aedes aegypti (L.) were compared using isozyme starch gel electrophoresis to characterize genetic variation between populations. Ae. aegypti were collected from seven provinces in Thailand. Thirty‐one isozyme encoding loci, including 19 polymorphic loci, were characterized. Only small levels of genetic differentiation were observed among the 31 district populations in the seven provinces. Isolation by distance among populations from the seven provinces showed no correlation between genetic variation and geographical distance.