Movement Behavior of Native and Invasive Small Mammals Shows Logging May Facilitate Invasion in a Tropical Rain Forest

Invasive species pose one of the greatest threats to biodiversity. This study investigates the extent to which human disturbance to natural ecosystems facilitates the spread of non‐native species, focusing on a small mammal community in selectively logged rain forest, Sabah, Borneo. The microhabitat preferences of the invasive Rattus rattus and three native species of small mammal were examined in three‐dimensional space by combining the spool‐and‐line technique with a novel method for quantifying fine‐scale habitat selection. These methods allowed the detection of significant differences for each species between the microhabitats used compared with alternative, available microhabitats that were avoided. Rattus rattus showed the greatest preference for heavily disturbed habitats, and in contrast to two native small mammals of the genus Maxomys, R. rattus showed high levels of arboreal behavior, frequently leaving the forest floor and traveling through the understory and midstory forest strata. This behavior may enable R. rattus to effectively utilize the complex three‐dimensional space of the lower strata in degraded forests, which is characterized by dense vegetation. The behavioral flexibility of R. rattus to operate in both terrestrial and arboreal space may facilitate its invasion into degraded forests. Human activities that generate heavily disturbed habitats preferred by R. rattus may promote the establishment of this invasive species in tropical forests in Borneo, and possibly elsewhere. We present this as an example of a synergistic effect, whereby forest disturbance directly threatens biodiversity and indirectly increases the threat posed by invasive species, creating habitat conditions that facilitate the establishment of non‐native fauna.     

Diversity and Coexistence of Ectoparasites in Small Rodents in a Tropical Dry Forest

To understand the mechanisms driving species diversity is central to community ecology. Here, we explored if habitat partitioning is associated with a species‐rich ectoparasite community in small rodents from a tropical dry forest in western Mexico. We trapped 199 mice in three 0.5 ha‐plots from eight small rodent species for every two months, from July 2011 to April 2012, and collected their ectoparasites. We identified 17 species of mites, two sucking lice species, two phoretic species, and one commensal species. The most abundant ectoparasite species was Steptolaelaps liomydis, representing 42 percent of all ectoparasites collected; seven ectoparasite species had < 10 individuals. Eighteen ectoparasite species (of 22 species) were collected from the most abundant rodent Liomys pictus. C‐score and the number of checkerboard species pairs were significantly higher against a random expectation. Ectoparasite species in L. pictus mice showed host microhabitat partitioning; Fahrenholzia ehrlichi and Fahrenholzia texana were found only in the anterior dorsal area, Ornithonysus sp. occurred along the dorsal part, Ixodes species were restricted to the ears, and Steptolaelaps liomydis was found throughout the body. We also identified ectoparasite communities with distinct species composition in two rodent species that use contrasting macrohabitats (L. pictus, strictly terrestrial; Peromyscus perfulvus, mostly arboreal). The remaining and low abundant rodent species showed a species‐poor ectoparasite community composition. We conclude that habitat partitioning at both macro and microhabitat scales appeared to characterize the species‐rich ectoparasite community. Conversely, most rodent host species with low abundances showed a species‐poor ectoparasite community.     

Microhabitat associations of terrestrial insectivorous birds in Amazonian rainforest and second‐growth forests

ABSTRACT Across the Neotropics, small‐bodied terrestrial insectivores are sensitive to forest fragmentation and are largely absent from second‐growth forests. Despite their sensitivity to forest structure, the microhabitat relationships of these birds have not been quantified. From July 1994 to January 1995 in central Amazonia, we characterized habitat at sites where nine species of terrestrial insectivores were observed foraging, as well as at randomly selected sites in continuous forest and two types of 10–15‐yr‐old second‐growth forest common in Amazonia (Vismia‐ and Cecropia‐dominated). We used factor analysis to find suites of correlated variables. From each factor, we selected a representative variable that was relatively easy to measure. We used Bayesian analysis to estimate means and standard deviations of these variables for each species and for each type of habitat. All nine focal species were associated with ranges of microhabitat variables, such as leaf litter depth and tree densities, often absent in second‐growth forests. At least in the early stages of regeneration, neither type of second‐growth forest provides suitable structure for the terrestrial insectivores in our study. The large leaves of Cecropia trees that make up the thick leaf litter may preclude the use of Cecropia‐dominated second growth by our focal species, many of which manipulate leaves when foraging. The leaf litter in Vismia‐dominated second growth was also thicker than sites used for foraging by our focal species. In addition, Vismia‐dominated growth had more small trees and small nonwoody vegetation, perhaps impeding movement by terrestrial birds. In continuous forest, our focal species foraged in microhabitats with characteristics that generally overlapped those of randomly selected sites. Thus, our results are consistent with the hypothesis that microhabitat differences make second‐growth forests unsuitable for our focal species.     

Habitat associations of opossums and rodents in a lowland forest in French Guiana

Opossums and rodents were sampled by live-trapping at Paracou, French Guiana to study habitat use. Tomahawk and Sherman traps were placed on the ground and at various heights in the subcanopy and canopy. We captured 10 species of opossums and seven species of rodents. We measured 14 microhabitat variables at each of 485 sampling stations. Presence or absence at a trap station of the three most abundant species of opossums (Didelphis albiventris, Didelphis marsupialis, and Micoureus demerarae) and two rodents (Oecomys rutilus and Proechimys cuvieri) were related by logistic regression analysis to the microhabitat variables. All five species were associated with at least one habitat variable. Linear analysis of categorical data was used to determine the degree of arboreality. Didelphis marsupialis and P. cuvieri were more commonly captured on the ground, D. albiventris was equitably distributed between ground and arboreal traps, and the remaining species were mostly arboreal. Linear analysis was used to compare the distribution of arboreal captures among two substrate types, two heights, and two substrate diameters. Oecomys rutilus was more commonly captured on lianas, D. albiventris and O. rutilus were more commonly captured at lower heights, and D. marsupialis demonstrated a preference for larger arboreal substrates.     

Topographic microclimates drive microhabitat associations at the range margin of a butterfly

The habitat associations of individuals underpin the dynamics of species distributions. Broad‐scale gradients in climate can alter habitat associations across species’ geographic ranges, but topographic heterogeneity creates local microclimates which could generate variation in habitat use at finer spatial scales. We examined the selection of microhabitats for egg‐laying by populations of a thermally‐constrained butterfly, the skipper Hesperia comma, across 16 sites with different regional temperatures and topographic microclimates. Using models of thermal microclimate, we examined how the association between eggs and warm bare ground microhabitats varied with ambient temperature, and predicted bare ground associations in 287 existing H. comma populations, to investigate the relative impacts of regional temperatures and topographic microclimates on microhabitat use. Eggs were most strongly associated with bare ground in relatively cool sites, indicating climate‐driven changes in microhabitat use. The majority of temperature variation between study sites was attributable to topographic microclimates rather than regional temperature differences, such that changes in microhabitat associations occurred principally between north‐ and south‐facing slopes within the same region. Predicted microhabitat associations across the UK distribution of H. comma showed that, due to the large temperature differences generated by topography, most of the between‐population variation in microhabitat use occurs locally within 5 km grid squares, with a smaller proportion occurring at a regional level between 5 km squares. Our findings show how microclimatic variation generated by topography alters the habitat associations of populations at fine spatial scales, suggesting that microclimate‐driven changes in habitat suitability could shape species’ distribution dynamics and their responses to environmental change.     

Feeding guild structure of beetles on Australian tropical rainforest trees reflects microhabitat resource availability

1.?We tested the hypotheses that feeding guild structure of beetle assemblages changed with different arboreal microhabitats and that these differences were consistent across rainforest tree species. 2.?Hand collection and beating techniques were used from the gondola of the Australian Canopy Crane to collect beetles from five microhabitats (mature leaves, flush leaves, flowers, fruit and suspended dead wood) within the rainforest canopy. A simple randomization procedure was implemented to test whether the abundances of each feeding guild on each microhabitat were different from that expected based on a null hypothesis of random distribution of individuals across microhabitats. 3.?Beetles from different feeding guilds were not randomly distributed, but congregated on those microhabitats that are likely to provide the highest concentrations of their preferred food sources. Herbivorous beetles, in particular, were over-represented on flowers and flush foliage and under-represented on mature leaves and dead wood. Proportional numbers of species within each feeding guild were remarkably uniform across tree species for each microhabitat, but proportional abundances of feeding guilds were all significantly non-uniformly distributed between host tree species, regardless of microhabitat, confirming patterns previously found for arthropods in trees in temperate and tropical forests. 4.?These results show that the canopy beetle community is partitioned into discrete assemblages between microhabitats and that this partitioning arises because of differences in feeding guild structure as a function of the diversity and the temporal and spatial availability of resources found on each microhabitat.     

Evolution of morphology and locomotor performance in anurans: relationships with microhabitat diversification

The relationships between morphology, performance, behavior and ecology provide evidence for multiple and complex phenotypic adaptations. The anuran body plan, for example, is evolutionarily conserved and shows clear specializations to jumping performance back at least to the early Jurassic. However, there are instances of more recent adaptation to habit diversity in the post‐cranial skeleton, including relative limb length. The present study tested adaptive models of morphological evolution in anurans associated with the diversity of microhabitat use (semi‐aquatic arboreal, fossorial, torrent, and terrestrial) in species of anuran amphibians from Brazil and Australia. We use phylogenetic comparative methods to determine which evolutionary models, including Brownian motion (BM) and Ornstein‐Uhlenbeck (OU) are consistent with morphological variation observed across anuran species. Furthermore, this study investigated the relationship of maximum distance jumped as a function of components of morphological variables and microhabitat use. We found there are multiple optima of limb lengths associated to different microhabitats with a trend of increasing hindlimbs in torrent, arboreal, semi‐aquatic whereas fossorial and terrestrial species evolve toward optima with shorter hindlimbs. Moreover, arboreal, semi‐aquatic and torrent anurans have higher jumping performance and longer hindlimbs, when compared to terrestrial and fossorial species. We corroborate the hypothesis that evolutionary modifications of overall limb morphology have been important in the diversification of locomotor performance along the anuran phylogeny. Such evolutionary changes converged in different phylogenetic groups adapted to similar microhabitat use in two different zoogeographical regions.     

Macroevolution of arboreality in salamanders

Evolutionary theory predicts that selection in distinct microhabitats generates correlations between morphological and ecological traits, and may increase both phenotypic and taxonomic diversity. However, some microhabitats exert unique selective pressures that act as a restraining force on macroevolutionary patterns of diversification. In this study, we use phylogenetic comparative methods to investigate the evolutionary outcomes of inhabiting the arboreal microhabitat in salamanders. We find that arboreality has independently evolved at least five times in Caudata and has arisen primarily from terrestrial ancestors. However, the rate of transition from arboreality back to terrestriality is 24 times higher than the converse. This suggests that macroevolutionary trends in microhabitat use tend toward terrestriality over arboreality, which influences the extent to which use of the arboreal microhabitat proliferates. Morphologically, we find no evidence for an arboreal phenotype in overall body proportions or in foot shape, as variation in both traits overlaps broadly with species that utilize different microhabitats. However, both body shape and foot shape display reduced rates of phenotypic evolution in arboreal taxa, and evidence of morphological convergence among arboreal lineages is observed. Taken together, these patterns suggest that arboreality has played a unique role in the evolution of this family, providing neither an evolutionary opportunity, nor an evolutionary dead end.     

Microhabitat Preference of Egernia napoleonis in Undisturbed Jarrah Forest,and Availability and Introduction of Microhabitats to Encourage Colonization of Restored Forest

An animal's microhabitat requirements can impact its ability to colonize restored areas, particularly species requiring slow developing microhabitats, such as logs and woody debris piles. Introduction of these microhabitats may be required to facilitate colonization by some species. Restored bauxite mine‐pits in the Jarrah (Eucalyptus marginata) forest of south‐western Australia contain introduced log piles at densities of 1 ha^?1. However, these have not facilitated colonization by Napoleon's skink (Egernia napoleonis), which rely on logs for habitat and are largely absent from restored sites. We radio‐tracked 12 skinks in unmined forest to determine their microhabitat preferences and examined differences in vegetation structure, and microhabitat and food availability, between restored and unmined forests to identify reasons for their absence. Restored and unmined forests differed in canopy, mid‐ and understory cover and ground substrates, which were all potential barriers to colonization. Food availability was similar between restored and unmined forest, thus not a barrier to colonization. Skinks primarily utilized long logs, large woody debris piles, and large trees; microhabitats that were scarce or absent in restored sites and, therefore, potential barriers to colonization. Using this information, we introduced small woody debris piles into restored sites in close proximity to unmined areas containing skinks to facilitate skink colonization. This showed early signs of success and suggested that the lack of logs and woody debris were barriers to colonization. However, further monitoring is required to accurately determine the long‐term value of woody debris piles in facilitating skink colonization.     

Generalist trophic ecology in a changing habitat: The case of the four‐striped mouse in a woody‐encroached savannah

Trophic generalists tolerate greater habitat change than specialists; however, few studies explore how generalist trophic ecology is affected. We established how the trophic ecology of an extreme generalist, Rhabdomys pumilio, changed in relation to a directionally changing woody‐encroached savannah in Eastern Cape, South Africa by investigating (a) foraging behaviour, (b) trophic niche and (c) feedback effects. (a) Giving‐up densities showed that R. pumilio preferred foraging in subcanopy microhabitat during the night as a result of lower thermoregulatory costs, but had similar preferences for sub‐ and intercanopy microhabitats during the day. (b) An isotope analysis revealed that the dietary composition and trophic niche occupied by R. pumilio differed among tree canopy cover levels (0%, 30% and 80%), which appeared to be related to changes in C₄ grass material and invertebrate availability. (c) Artificial seed patches suggested that R. pumilio was a potentially important postdispersal seed predator of the woody‐encroaching species, Vachellia karroo. Thus, an increase in tree canopy cover altered the trophic niche of R. pumilio by reducing foraging costs at night and providing alternative food resources in terms of availability and source. These findings demonstrate how an extreme generalist adapted to human‐induced habitat change through changes in its trophic ecology.     

Effects of tree species diversity on a community of weaver spiders in a tropical forest plantation

The effects of producer diversity on predators have received little attention in arboreal plant communities, particularly in the tropics. This is particularly true in the case of tree diversity effects on web‐building spiders, one of the most important groups of invertebrate predators in terrestrial plant communities. We evaluated the effects of tree species diversity on the community of weaver spiders associated with big‐leaf mahogany (Swietenia macrophylla) in 19, 21 × 21‐m plots (64 plants/plot) of a tropical forest plantation which were either mahogany monocultures (12 plots) or polycultures (seven plots) that included mahogany and three other tree species. We conducted two surveys of weaver spiders on mahogany trees to evaluate the effects of tree diversity on spider abundance, species richness, diversity, and species composition associated with mahogany. Our results indicated that tree species mixtures exhibited significantly greater spider abundance, species richness, and diversity, as well as differences in spider species composition relative to monocultures. These results could be due to species polycultures providing a broader range of microhabitat conditions favoring spider species with different habitat requirements, a greater availability of web‐building sites, or due to increased diversity or abundance of prey. Accordingly, these results emphasize the importance of mixed forest plantations for boosting predator abundance and diversity and potentially enhancing herbivore pest suppression. Future work is necessary to determine the specific mechanisms underlying these patterns as well as the top‐down effects of increased spider abundance and species richness on herbivore abundance and damage.     

Vertical distribution of Hymenophyllaceae species among host tree microhabitats in a temperate rain forest in Southern Chile

Question: Are differences in microhabitat preferences of co‐occurring epiphytic Hymenophyllaceae species (filmy ferns) correlated with differences in ecophysiological responses to light availability and humidity in the host tree? Location: The Andean foothills in south‐central Chile. Methods: We evaluated the distribution pattern of nine filmy fern species in microhabitats that differ in light availability and humidity in four host tree species. A DCA was developed to assess Hymenophyllaceae species microhabitat preference in terms of canopy openness (CO) and relative humidity. We assessed whether differences in chlorophyll content, maximum photochemical efficiency (Fv/Fm), photosynthetic capacity (A_max), evapotranspiration (E) and instantaneous water use efficiency (WUE) are consistent with any pattern. Results: CO and relative humidity differed significantly with height in the host trees. While CO increased with height in a host tree, relative humidity decreased. DCA analysis showed that filmy fern species distribution within and among trees was mainly explained by the relative humidity of the microhabitat. Chlorophyll content, chlorophyll a/b ratio, A_max and E differed significantly among filmy fern species. A_max and E were correlated with axis 1 scores from the DCA analysis. Conclusions: The vertical distribution and abundance of filmy fern species in Chilean temperate rain forest seems to be closely related to the different microhabitats offered by host trees. This pattern may reflect interspecific differences in ecophysiological traits related both to light availability and humidity. Our results suggest that humidity is the main environmental factor driving functional responses and habitat preferences of these filmy fern species.     

Intraguild dynamics of understudied carnivores in a human‐altered landscape

Mesocarnivores constitute a diverse and often abundant group of species, which are increasingly occupying hweigher trophic levels within multi‐use landscapes. Yet, we know relatively little about their interactions with each other, especially in human‐altered areas. Using camera trap data collected in a forestry concession in the Greater Gorongosa ecosystem of central Mozambique, we examined the spatiotemporal relationships and potential for intraguild competition among three understudied African carnivores: African civets (Civettictis civetta), bushy‐tailed mongooses (Bdeogale crassicauda), and large‐spotted genets (Genetta maculata). After accounting for habitat preferences and tolerance to anthropogenic factors, we found that African civets and bushy‐tailed mongooses avoid each other spatially and temporally. Additionally, civets and mongooses were also both more likely to use sites farther away from human settlements, possibly decreasing the total available habitat for each species if competition is driving this spatial partitioning. In contrast, we did not find evidence for spatial or temporal partitioning between large‐spotted genets and African civets, but bushy‐tailed mongooses altered their activity patterns where they co‐occurred with genets. Our study contributes to scant ecological knowledge of these mesocarnivores and adds to our understanding of community dynamics in human‐altered ecosystems.     

Aspects of community ecology of amphibians and reptiles at Bonny Island (Nigeria), an area of priority relevance for petrochemical industry

Finima Nature Park, situated in Bonny Island (Niger Delta, Nigeria), is a protected forested area placed within one of the most industrially developed (and environmentally polluted) regions of the whole African continent. Amphibian and reptile community composition in relation to season and microhabitat characteristics was studied, by a combination of field techniques, during the wet and dry seasons of 2007 and 2008. Overall, a total of 21 species of reptiles from nine families (with 668 individuals caught) and eight amphibian species from five families (492 individuals caught) were recorded. Nearly 94% of the amphibian individuals and most of the species were recorded, especially during the wet season, whereas most of the reptiles were found with no inter‐seasonal differences. However, Agama agama and Mabuya affinis were found especially during the dry season. As for microhabitat use, the reptile species showed a clear, aggregated preference for two microhabitat types (dry leaves and under logs), which are probably crucial when these animals need to lose body temperature to avoid overheating. Concerning amphibians, the various species used the available microhabitats in a way expectable from their main ecological traits, with toads differing from aquatic frogs, and these differing from arboreal frogs.     

Contrasting habitat use of morphologically similar bat species with differing conservation status in south‐eastern Australia

The east‐coast free‐tailed bat Mormopterus norfolkensis Gray, 1839 is a threatened insectivorous bat that is poorly known and as such conservation management strategies are only broadly prescribed. Insectivorous bats that use human‐modified landscapes are often adapted to foraging in open microhabitats. However, few studies have explored whether open‐adapted bats select landscapes with more of these microhabitat features. We compared three morphologically similar and sympatric, molossid bats (genus Mormopterus) with different conservation status in terms of their association with vegetation, climate, landform and land‐use attributes at landscape and local habitat element scales. We predicted that these species would use similar landscape types, with semi‐cleared and low density urban landscapes used more than forested and heavily cleared landscapes. Additionally, we explored which environmental variables best explained the occurrence of each species by constructing post‐hoc models and habitat suitability maps. Contrary to predictions, we found that the three species varied in their habitat use with no one landscape type used more extensively than other types. Overall, M. norfolkensis was more likely to occur in low‐lying, non‐urban, riparian habitats with little vegetation cover. Mormopterus species 2 occupied similar habitats, but was more tolerant of urban landscapes. In contrast, Mormopterus species 4 occurred more often in cleared than forested landscapes, particularly dry landscapes with little vegetation cover. The extensive use of coastal floodplains by the threatened M. norfolkensis is significant because these habitats are under increasing pressure from human land‐uses and the predicted increase in urbanization is likely to further reduce the amount of suitable habitat.     

Correlates of active body temperatures and microhabitat occupation in nine species of central Australian agamid lizards

Body temperatures of active lizards and their correspondence with microhabitat occupation were studied for nine species of agamid lizards in the central Australian arid zone. Thermoregulatory behaviour was also documented using several measures, such as the use of shade and perch height. The effects of thermal environment on lizard habitat occupation were hypothesized to be significant, because desert regions experience daily and seasonal extremes of temperature that are well in excess of a lizard’s preferred temperature range. All species, except Ctenophorus isolepis and Diporiphora winneckei, were found to have body temperatures that corresponded closely to ground and surface temperatures. Thermoregulatory behaviour was also found to be important throughout a lizard’s daily activity; all study species, other than Ctenophorus isolepis, were found to increase their perch height in the middle of the day. Ctenophorus isolepis was shown to be a strictly terrestrial species that uses the shade of spinifex in its thermoregulatory behaviour. Species exhibited a non‐random selection of microhabitats and a preference for a particular set of thermal and structural factors. In this study, it was shown that structural factors were particularly important in microhabitat occupation. Thermal factors accounted for a smaller proportion of variance in microhabitat occupation, but still played a considerable role in the microhabitat use in central Australian agamids.     

Effects of temperature and elevation on habitat use by a rare mountain butterfly: implications for species responses to climate change

Abstract 1. The present study used the mountain specialist butterfly Parnassius apollo as a model system to investigate how climate change may alter habitat requirements for species at their warm range margins. 2. Larval habitat use was recorded in six P. apollo populations over a 700 m elevation gradient in the Sierra de Guadarrama (central Spain). Larvae used four potential host species (Sedum spp.) growing in open areas amongst shrubs. 3. Parnassius apollo host‐plant and habitat use changed as elevation increased: the primary host shifted from Sedum amplexicaule to Sedum brevifolium, and larvae selected more open microhabitats (increased bare ground and dead vegetation, reduced vegetation height and shrub cover), suggesting that hotter microhabitats are used in cooler environments. 4. Larval microhabitat selection was significantly related to ambient temperature. At temperatures lower than 27 °C, larvae occupied open microhabitats that were warmer than ambient temperature, versus more shaded microhabitats that were cooler than ambient conditions when temperature was higher than 27 °C. 5. Elevational changes in phenology influenced the temperatures experienced by larvae, and could affect local host‐plant favourability. 6. Habitat heterogeneity appears to play an important role in P. apollo larval thermoregulation, and may become increasingly important in buffering populations of this and other insect species against climatic variation.