Heterothermy in Afrotropical mammals and birds: a review

Recent years have seen a rapid increase in the number of Afrotropical endotherms known to avoid mismatches between energy supply and demand by using daily torpor and/or hibernation. Among mammals, heterothermy has been reported in 40 species in six orders, namely Macroscelidea, Afrosoricida, Rodentia, Eulipotyphla, Primates and Chiroptera. These species span a range in body mass of 7-770?g, with minimum heterothermic body temperatures ranging from 1-27°C and bout length varying from 1?h to 70 days. Daily torpor is the most common form of heterothermy, with true hibernation being observed in only seven species, Graphiurus murinus, Graphiurus ocularis, Atelerix frontalis, Cheirogaleus medius, Cheirogaleus major, Microcebus murinus and Microcebus griseorufus. The traditional distinction between daily torpor and hibernation is blurred in some species, with free-ranging individuals exhibiting bouts of > 24?h and body temperatures < 16 °C, but none of the classical behaviours associated with hibernation. Several species bask in the sun during rewarming. Among birds, heterothermy has been reported in 16 species in seven orders, and is more pronounced in phylogenetically older taxa. Both in mammals and birds, patterns of heterothermy can vary dramatically among species occurring at a particular site, and even among individuals of a single species. For instance, patterns of heterothermy among cheirogalid primates in western Madagascar vary from daily torpor to uninterrupted hibernation for up to seven months. Other examples of variation among closely-related species involve small owls, elephant shrews and vespertilionid bats. There may also be variation in terms of the ecological correlates of torpor within a species, as is the case in the Freckled Nightjar Caprimulgus tristigma.     

Relationship Between Microhabitat Structure and Distribution of Mouse Lemurs (Microcebus spp.) in Northwestern Madagascar

Understanding processes affecting the distribution and abundance of organisms is a central issue in ecology and conservation biology. In northwestern Madagascar, we found an uneven distribution pattern of the golden-brown mouse lemur (Microcebus ravelobensis) and the grey mouse lemur (M. murinus). In one area (JBA) the two species lived sympatrically, whereas in another forest area (JBB) Microcebus ravelobensis occurred exclusively. To investigate whether differences in forest structure may explain this uneven distribution, we conducted a microhabitat analysis and related it to specific distribution. In JBA the habitat of Microcebus ravelobensis was characterized by a higher percentage of trees with many lianas and a higher cover of the herb layer, whereas that of M. murinus had a higher number of trees with a diameter at breast height (DBH) >10 cm. The comparison of the forest structure of the microhabitats of the two species between JBA and JBB revealed further differences. The cover of the overstory, the percentage of trees without lianas and the number of larger trees (DBH >10 cm) among the microhabitats were higher for Microcebus murinus in JBA than for M. ravelobensis in JBB whereas the microhabitats of M. ravelobensis at the two sites did not differ concerning these vegetation structures. Differences between the two species coincide with those of resources important for survival. Our results indicate the importance of microhabitat analyses for the understanding of distribution patterns of species and for successful conservation planning.     

Habitat use and diet of two stream gobies of the genus Rhinogobius in south-western Shikoku,Japan

The importance of interspecific competition to habitat use by two congeneric stream gobies, Rhinogobius sp. LD (large-dark type) and CB (cross-band type), was studied by: (i) examining differences in habitat use by each species along the course of the stream; and (ii) comparing microhabitat use and the diet of each species between in allopatry and in sympatry in tributaries of the Shimanto River, south-western Shikoku, Japan. Rhinogobius sp. LD mainly used riffles through the course of the stream. Although CB also mainly used riffles in the lower reaches where CB was numerically dominant, the greater use of riffles by CB was not observed in upper reaches where LD was dominant. Microhabitat analysis revealed that both LD and CB preferred lower bottom velocity in faster current in both allopatry and sympatry. In sympatry, however, LD used coarser substrate and faster current than CB, and both LD and CB used narrower ranges of microhabitat conditions than in allopatry. Dietary analysis indicated that both allopatric and sympatric LD preferred baetids, which are apt to drift, suggesting they adopted ambush foraging. Allopatric CB preferred leptophlebiids and chironomids, which are also apt to drift, whereas sympatric CB did not prefer such invertebrates. Overall results of this study suggest that LD and CB compete for better feeding habitats, CB shift their habitat use and foraging mode under the influence of LD, and current velocity and substrate coarseness are key factors in their habitat segregation.

     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Sharing the Space: Distribution,Habitat Segregation and Delimitation of a New Sympatric Area of Subterranean Rodents

Subterranean rodents of the genus Ctenomys usually present an allopatric or parapatric distribution. Currently, two cases of sympatry have been recognized for the genus in the coastal dunes of southern Argentina and southern Brazil. In this context, they are ideal models to test hypotheses about the factors that delimit the patterns of space use and to understand interspecific interactions in small mammals. We investigated the vegetation structure, plant biomass and soil hardness selected by two species of subterranean rodents (Ctenomys flamarioni and C. minutus) when distributed in sympatry and allopatry from nine different areas along the line of coastal dunes in southern Brazil. In addition, our work presents a new record of a third area of sympatry for the genus Ctenomys. Ctenomys flamarioni and C. minutus show habitat segregation in the area where they occur in sympatry. These species show segregation in their selection of microhabitats, differing in relation to soil hardness, plant biomass, and plant cover. Ctenomys flamarioni showed a distinction in habitat selection when occurring in allopatry and sympatry, whereas C. minutus selected the same habitat characteristics under both conditions. A possible explanation to the observed pattern is that these species have acquired different adaptations over time which allows them the ability to exploit different resources and thus avoid competitive interactions all together.     

Adaptive loss of color polymorphism and character displacements in sympatric <Emphasis Type="Italic">Mnais</Emphasis> damselflies

A geographical survey of two Mnais damselfly species in the Kinki area of Japan showed evidence for character displacements when the two species were found in sympatry. Mnais costalis, a species that has polymorphic male mating types of orange-winged territorial and clear-winged non-territorial morphs (hereafter abbreviated to orange and clear morphs respectively) in allopatry often shifted to having monomorphic orange morphs in sympatry. The mean body size of orange morphs was consistently larger than that of clear morph in allopatry. The mean body size of the sympatric orange morphs was even larger than that of allopatric orange morphs. By contrast, Mnais pruinosa, a species that also has two morphs of large orange and small clear morphs in allopatry, shifted to having monomorphic clear morphs in sympatry. The mean body size of the sympatric clear morphs was smaller than that of allopatric clear morphs. Divergence was also detected in the preference for habitat insolation conditions between sympatric Mnais damselflies. Both species in allopatric regions prefer half-light forest habitats, while in sympatric regions they showed diversified habitat preference: M. costalis preferred sunny habitats while M. pruinosa preferred shady habitats. Multiple character displacements in signal traits and habitat preference emerged in heterogeneous forest light environments are likely to have synergistic effects on the reproductive isolation of the two species.     

Microhabitat distribution of two Florida scrub endemic plants in comparison to their habitat-generalist congeners

Habitat-specialist species may be restricted to a narrower range of microhabitats than habitat-generalist species. We addressed this hypothesis by comparing microhabitats of two pairs of congeners that differ in habitat specificity and co-occur in one distinct habitat type, Florida rosemary scrub. We characterized microhabitats of rosemary scrub specialists, Polygonella basiramia and Lechea cernua, their habitat-generalist congeners, Polygonella robusta and Lechea deckertii, and random points in the rosemary scrub habitat. Plants of both habitat specialists occurred in microhabitats with significantly more bare sand than plants of habitat-generalist species and random points. Plants of all four species occurred in microhabitats that were farther from dominant shrubs, Ceratiola and Quercus spp., than random points. Seedlings of both habitat specialists grew larger in bare sand microhabitats, whereas ground lichens and litter did not affect seedling growth of the habitat generalists. As the time since fire increases, bare sand cover decreases, Ceratiola density increases, Quercus density remains constant, and shrubs become taller. Physical characteristics, such as soil temperature, soil carbon, and soil moisture, differ slightly with respect to microhabitat. Our results suggest that P. basiramia and L. cernua are specialized on bare sand microhabitats that characterize their preferred habitat, rosemary scrub. Microhabitat specialization may limit the distribution of these rare species.     

Effect of competition on habitat utilization in two temperate climate gecko species

Competition over spatial niche utilisation is one of most common competitive interactions between species in sympatry. Moreover, competitive interactions may involve age classes, and can fluctuate temporally. Consequently, evasive strategies that enable co-existence are likely to be important in the evolution of species assemblages. Here we investigate a system of two co-existing species of temperate geckos with similar ecologies (the house gecko, Hemidactylus turcicus and the wall gecko, Tarentola mauritanica), providing an opportunity to study the effect of species interactions. Juveniles and adults of both species were investigated throughout their daily and annual cycle to explore the effect of inter- and intra-specific interactions on microhabitat use. The two species showed differences in habitat use for both age classes in sympatry. In sympatry, T. mauritanica uses more open habitats and is more active. In contrast, H. turcicus is found in more closed habitats, closer to the ground and to vegetation cover. In allopatry, H. turcicus was observed in more open habitats, closer to the ground, and to vegetation cover, when compared to the population in sympatry with T. mauritanica. These differences in habitat usage were significant for both age classes. Moreover, there were differences, both in sympatry and in allopatry, between age classes that were dependent on season. In conclusion, the presence of a competitor induces a spatial shift in individuals of both age classes of H. turcicus. Observed plasticity in habitat utilisation in both age classes of H. turcicus is used to argue for the invasive potential of this species.     

Space Use Variation in Co-Occurring Sister Species: Response to Environmental Variation or Competition?

Coexistence often involves niche differentiation either as the result of environmental divergence, or in response to competition. Disentangling the causes of such divergence requires that environmental variation across space is taken into account, which is rarely done in empirical studies. We address the role of environmental variation versus competition in coexistence between two rodent species: Rhabdomys bechuanae (bechuanae) and Rhabdomys dilectus dilectus (dilectus) comparing their habitat preference and home range (HR) size in areas with similar climates, where their distributions abut (allopatry) or overlap (sympatry). Using Outlying Mean Index analyses, we test whether habitat characteristics of the species deviate significantly from a random sample of available habitats. In allopatry, results suggest habitat selection: dilectus preferring grasslands with little bare soil while bechuanae occurring in open shrublands. In sympatry, shrubland type habitats dominate and differences are less marked, yet dilectus selects habitats with more cover than bechuanae. Interestingly, bechuanae shows larger HRs than dilectus, and both species display larger HRs in sympatry. Further, HR overlaps between species are lower than expected. We discuss our results in light of data on the phylogeography of the genus and propose that evolution in allopatry resulted in adaptation leading to different habitat preferences, even at their distribution margins, a divergence expected to facilitate coexistence. However, since sympatry occurs in sites where environmental characteristics do not allow complete species separation, competition may explain reduced inter-species overlap and character displacement in HR size. This study reveals that both environmental variation and competition may shape species coexistence.     

Why Do Species Co-Occur? A Test of Alternative Hypotheses Describing Abiotic Differences in Sympatry versus Allopatry Using Spadefoot Toads

Areas of co-occurrence between two species (sympatry) are often thought to arise in regions where abiotic conditions are conducive to both species and are therefore intermediate between regions where either species occurs alone (allopatry). Depending on historical factors or interactions between species, however, sympatry might not differ from allopatry, or, alternatively, sympatry might actually be more extreme in abiotic conditions relative to allopatry. Here, we evaluate these three hypothesized patterns for how sympatry compares to allopatry in abiotic conditions. We use two species of congeneric spadefoot toads, Spea multiplicata and S. bombifrons, as our study system. To test these hypotheses, we created ecological niche models (specifically using Maxent) for both species to create a map of the joint probability of occurrence of both species. Using the results of these models, we identified three types of locations: two where either species was predicted to occur alone (i.e., allopatry for S. multiplicata and allopatry for S. bombifrons) and one where both species were predicted to co-occur (i.e., sympatry). We then compared the abiotic environment between these three location types and found that sympatry was significantly hotter and drier than the allopatric regions. Thus, sympatry was not intermediate between the alternative allopatric sites. Instead, sympatry occurred at one extreme of the conditions occupied by both species. We hypothesize that biotic interactions in these extreme environments facilitate co-occurrence. Specifically, hybridization between S. bombifrons females and S. multiplicata males may facilitate co-occurrence by decreasing development time of tadpoles. Additionally, the presence of alternative food resources in more extreme conditions may preclude competitive exclusion of one species by the other. This work has implications for predicting how interacting species will respond to climate change, because species interactions may facilitate survival in extreme habitats.     

Different Population Dynamics of Microcebus murinus in Primary and Secondary Deciduous Dry Forests of Madagascar

The goal of this study was to identify causes for lower population densities of mouse lemurs (Microcebus murinus) in secondary than in primary dry deciduous forests of western Madagascar. Variations in the abundance of Microcebus murinus are linked to the capacity to enter energy-saving torpor during the dry season. Under natural conditions in primary forest, Microcebus murinus can maintain daily torpor (and possibly hibernation) as long as body temperatures remain below 28°C. Females are more likely to hibernate than males, resulting in skewed sex ratios of captured Microcebus murinus in the primary forest. In the secondary forest the sex ratio of subjects captured during the dry season is more balanced than in primary forest, indicating that fewer females go into torpor in secondary than in primary forest. Secondary forests have fewer large standing or fallen trees that might provide holes as shelter for Microcebus murinus. Ambient temperatures are higher in secondary than in primary forests and higher outside than inside tree holes. These high ambient temperatures might hinder the ability of Microcebus murinus to maintain torpor for prolonged periods in secondary forests. Mouse lemurs from secondary forest have lower body mass than in primary forest. The year-to-year recapture rate is zero in secondary forest and thus significantly lower than in primary forest. This indicates that survival rates are lower in secondary than in primary forests. Thus, secondary forests may be of limited value as buffer zones or even corridors for mouse lemurs.     

Non‐native trout limit native brook trout access to space and thermal refugia in a restored large‐river system

We used direct observation via snorkeling surveys to quantify microhabitat use by native brook (Salvelinus fontinalis) and non‐native brown (Salmo trutta) and rainbow (Onchorynchus mykiss) trout occupying natural and restored pool habitats within a large, high‐elevation Appalachian river, United States. Permutational multivariate analysis of variance (PERMANOVA) and subsequent two‐way analysis of variance (ANOVA) indicated a significant difference in microhabitat use by brook and non‐native trout within restored pools. We also detected a significant difference in microhabitat use by brook trout occupying pools in allopatry versus those occupying pools in sympatry with non‐native trout—a pattern that appears to be modulated by size. Smaller brook trout often occupied pools in the absence of non‐native species, where they used shallower and faster focal habitats. Larger brook trout occupied pools with, and utilized similar focal habitats (i.e. deeper, slower velocity) as, non‐native trout. Non‐native trout consistently occupied more thermally suitable microhabitats closer to cover as compared to brook trout, including the use of thermal refugia (i.e. ambient–focal temperature >2°C). These results suggest that non‐native trout influence brook trout use of restored habitats by: (1) displacing smaller brook trout from restored pools, and (2) displacing small and large brook trout from optimal microhabitats (cooler, deeper, and lower velocity). Consequently, benefits of habitat restoration in large rivers may only be fully realized by brook trout in the absence of non‐native species. Future research within this and other large river systems should characterize brook trout response to stream restoration following removal of non‐native species.     

Microhabitat use in a mediterranean riverine fish assemblage

Summary We examined microhabitat use in Barbus graellsii, Barbus haasi, Chondrostoma toxostoma, Rutilus arcasii, and Salmo gairdneri over a 19 month period in the upper Rio Matarraña, Spain. B. graellsii and Ch. toxostoma exhibited non-random microhabitat use during all seasons and preferentially occupied deep microhabitats with heterogeneous substrates. During the majority of seasons in which they were present, B. haasi and R. arcasii occurred in microhabitats similar to those occupied by B. graellsii and Ch. toxostoma. S. gairdneri was over-represented in high velocity microhabitats with erosional substrates. We did not observe any evidence of interspecific interference competition or avoidance. Substrate composition did not appear to affect microhabitat use outside of its covariation with depth and velocity. Seaonal variation in microhabitat use by B. graellsii, B. haasi and Ch. toxostoma was strongly correlated with seasonal changes in microhabitat availability. S. gairdneri, however, occurred closer to the substrate when average velocities were high. Larger B. graellsii and B. haasi sometimes occupied deeper, higher velocity microhabitats than did smaller specimens. Larger B. graellsii also occasionally occurred farther from shelter than did smaller specimens; the reverse was true for B. haasi. Larger Ch. toxostoma sometimes were found farther from both the substrate and shelter than smaller individuals, whereas smaller specimens occasionally inhabited deeper areas with more depositional substrates than did larger Ch. toxostoma. During Late Summer 1985, smaller Ch. toxostoma also occupied microhabitats with higher velocities than did larger specimens. A comparison of microhabitat use for two species present in both upper and lower portions of the Matarraña indicated that most differences in microhabitat use could be attributed to inter-site differences in microhabitat availability. The data suggest, hovever, that both species shifted to more protected microhabitats in the higher velocity site. Assemblage members generally occupied statistically distinguishable microhabitats and could be classified as: 1) high-velocity upper water column (S. gairdneri), 2) low velocity lower water column (B. graellsii, Ch. toxostoma and R. arcasii), and 3) shelter-oriented benthic (B. haasi). The introduction of S. gairdneri during Winter 1984 did not produce microhabitat shifts in any of the native species. Whether or not the native species affected microhabitat use in S. gairdneri is unknown. Interspecific competition for space, however, did not appear to strongly influence microhabitat use among the native species.     

Reproductive character displacement in the acoustic communication system of green tree frogs (Hyla cinerea)

Abstract Interactions between species can affect the evolution of their sexual signals, receiver selectivity, or both. One commonly expected outcome is reproductive character displacement, whereby adverse consequences of mismating select for greater differentiation of communication systems in areas of sympatry than in areas of allopatry. We found evidence of reproductive character displacement in the acoustic communication system of green tree frogs ( Hyla cinerea ). The strength of female preferences for the spectral properties of calls that distinguish conspecific calls from those of a closely related congener, H. gratiosa , was greater in areas of sympatry with H. gratiosa than in areas of allopatry. We also found subtle differences in advertisement calls and in the heights of male calling perches when we restricted our comparisons to localities in which H. gratiosa was also breeding (syntopy) with localities where this species was absent. Hyla cinerea and H. gratiosa show only weak genetic incompatibility, but the calls representative of interspecific hybrids were unattractive to females of both parental species. Hybrids might also be at an ecological disadvantage because of different habitat preferences of the two taxa. Thus, selection against production of less fit or less attractive hybrid or backcross offspring are probably the main causes responsible for the differences documented in this paper.     

Changes in use of lake habitat by experimentally segregated populations of cutthroat trout and Dolly Varden char

The role of competition in a lacustrine community of two salmonid species, cutthroat trout Oncorhynchus clarki and Dolly Varden char Salvelinus malma , was studied in three coastal British Columbia lakes Habitat use by the species alone (allopatric) and in coexistence with each other (sympatric) was investigated by gill netting at 0-40 m depth contours (surface to bottom) so that several habitats (littoral, epipelagic. pelagic, epibenthic) were sampled From June to October, trout used mainly littoral and epipelagic habitats in sympatry and allopatry Char used all habitats in allopatry. and deep habitats (pelagic, epibenthic) not frequented by trout in sympatry The two species were thus spatially segregated with depth in sympatry Diel (day, night) and seasonal changes (spring, summer, autumn) in habitat use were not pronounced The shift in habitat use by experimentally allopatric char but not trout suggests that the effects of competition between sympatric trout and char for habitat resources are greater on the char     

Microhabitat preference constrains invasive spread of non-native natal grass (<Emphasis Type="Italic">Melinis repens</Emphasis>)

Preventing the establishment of a non-native species is critical for ensuring the species does not become invasive, yet most non-native species will have little impact on their environment. Despite this, little is known about what influences whether a species will remain relatively benign, or whether it will cause economic or ecological harm. Understanding a plant’s microhabitat provides insight into the necessary conditions for establishment and the current distribution limitations of a population. We investigated microhabitat preference of the non-native natal grass (Melinis repens (Willd.) Zizka) in Florida scrub using microhabitat sampling to measure vegetation composition. We examined the extent to which microhabitats were associated with natal grass presence and biomass in invaded disturbed scrub and roadside plots using backwards stepwise logistic regression and general linear models to identify significant microhabitat variables. We further compared these plots with those in undisturbed, uninvaded scrub to characterize vegetation across habitat types, and used our model to predict the probability of natal grass invasion in undisturbed scrub. Natal grass preferred microhabitats with high litter volume and distance to shrubs and intermediate cactus, graminoid, and vine cover. Roadside natal grass achieved higher biomass and was less microhabitat limited than disturbed scrub natal grass. We determined that undisturbed scrub plots represent distinct microhabitats that natal grass is unlikely to invade. Microhabitat sampling provides land-managers a non-intrusive technique to assess potential habitat suitability based non-native plant preferences before a costly invasion occurs.     

Habitat overlap between sympatric European hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) and Eastern cottontails (<Emphasis Type="Italic">Sylvilagus floridanus</Emphasis>) in northern Italy

In northern Italy, the range of the Eastern cottontail (Sylvilagus floridanus) largely overlaps with that of the native European hare (Lepus europaeus) on the Po Plain. Both species appear to have similar habitat requirements. We studied habitat selection by hares and cottontails during feeding activity from September 2006 to August 2007 in two areas where they occur alone (allopatry) and in one area where they occur together (sympatry). The three areas were basically similar, so that shifts in habitat use observed in sympatry should reflect the response to interspecific competition. Habitat selection was examined at micro- and macro-habitat levels throughout seasons. Habitat breadth of both species followed the change of resource availability through seasons in allopatry as well as in sympatry. No shifts in habitat use were evident at macro-habitat level, even during autumn which was the limiting season. Exploitation of shared habitats by the two species seems to be promoted by differential micro-habitat use within macro-habitat types. Cottontails used woods with dense understory in greater proportion than hares, and their present sites were concentrated within the maximum distance of 20 m of the nearest shelter site. Hares were more likely than cottontails to exploit crops, and their sites were distributed even greater than 80 m away from permanent cover patches. The habitat heterogeneity of agricultural ecosystems within the sympatry range could buffer the negative effects of external factors (climate, human disturbance and predation) on hares, and enhance the chances of exploitation of shared habitats by both species.     

Niche-habitat mechanisms and biotic interactions explain the coexistence and abundance of congeneric sandgrouse species

Ascertaining which niche processes allow coexistence between closely related species is of special interest in ecology. We quantified variations in the environmental niches and densities of two congeneric species, the pin-tailed and the black-bellied sandgrouse (Pterocles alchata and Pterocles orientalis) in allopatry and sympatry under similar abiotic, habitat and dispersal contexts to understand their coexistence. Using principal component analysis, we defined environmental gradients (niche dimensions) including abiotic, habitat and anthropogenic variables, and calculated niche breadth, position and overlap of both species in sympatry and allopatry. Additionally, sandgrouse density was modelled as a function of the niche dimensions and the density of the other species. We found evidence that each species occupies distinct environmental niches in sympatry and in allopatry. The black-bellied sandgrouse exploits a broader range of environmental conditions (wider niche breadth) while the pin-tailed sandgrouse reaches high densities where conditions seem to match its optimum. In sympatry, both species shift their niches to intermediate positions, indicating the importance of abiotic factors in setting coexistence areas. Environmental conditions determine regional densities of pin-tailed sandgrouse whereas biotic interactions explain the density of the black-bellied sandgrouse in areas with abiotic conditions similarly conducive for both species. Highly suitable areas for the pin-tailed sandgrouse fall beyond the upper thermal limit of the black-bellied sandgrouse, leading to niche segregation and low densities for the latter. Finally, local niche shift and expansion plus possible heterospecific aggregation allow the pin-tailed sandgrouse to thrive in a priori less favourable environments. This work provides insight into how different mechanisms allow species coexistence and how species densities vary in sympatry compared to allopatry as a result of environmental filtering and biotic interactions.     

Geographic variation in mimetic precision among different species of coral snake mimics

Batesian mimicry is widespread, but whether and why different species of mimics vary geographically in resemblance to their model is unclear. We characterized geographic variation in mimetic precision among four Batesian mimics of coral snakes. Each mimic occurs where its model is abundant (i.e. in ‘deep sympatry’), rare (i.e. at the sympatry/allopatry boundary or ‘edge sympatry’) and absent (i.e. in allopatry). Geographic variation in mimetic precision was qualitatively different among these mimics. In one mimic, the most precise individuals occurred in edge sympatry; in another, they occurred in deep sympatry; in the third, they occurred in allopatry; and in the fourth, precise mimics were not concentrated anywhere throughout their range. Mimicry was less precise in allopatry than in sympatry in only two mimics. We present several nonmutually exclusive hypotheses for these patterns. Generally, examining geographic variation in mimetic precision – within and among different mimics – offers novel insights into the causes and consequences of mimicry.