Vertical habitat segregation as a mechanism for coexistence in sympatric rodents

Coexistence has been widely studied in small mammals and frequently is assumed to be facilitated by habitat segregation. Using live trapping and spool-and-line experiments, we analyzed habitat selection and segregation across multiples scales for Peromyscus leucopus and Ochrotomys nuttalli. At the habitat scale of a forest stand (∼1–100s ha) P. leucopus co-occurred at all sites where O. nuttalli was found, and we did not detect evidence of positive or negative associations or habitat segregation. However, O. nuttalli was restricted to early successional forests, and P. leucopus had significantly lower abundances in early successional forests than in other habitats. We found similar patterns at the mesohabitat scale of the study site (400 m²). O. nuttalli abundance increased with increasing shrub and tree densities, while increases in P. leucopus abundance were associated with open understories. At the microhabitat scale of the individual movement trail, we found vertical segregation. Movement trails for O. nuttalli were at significantly higher elevation (mean height = 142.93 ± 37.10 cm) than P. leucopus trails (mean height = 15.4 ± 4.98 cm; F-value = 35.29, p < 0.001). We concluded that microhabitat segregation was driven by differential use of vertical space for movement and foraging. We suggest O. nuttalli is superior to P. leucopus in its ability to acquire food especially in shrubby subcanopies where few acorns are available. However, P. leucopus is superior in its ability to forage and avoid predators.     

Habitat utilization patterns of sympatric populations of Pseudomys gracilicaudatus and Rattus lutreolus in coastal heathland: A multivariate analysis

Abstract Habitat usage characteristics of two species of native murid rodents, Pseudomys gracilicaudatus and Rattus lutreolus were investigated on an area of coastal heathland at Myall Lakes National Park. A grid of 151 trap stations comprising 17 traplines was positioned across a mosaic of habitats. At each trap station 19 structural vegetation and physical variables known to affect the microdistribution of small mammals were measured. Multivariate statistical procedures identified those microhabitat variables that contribute to individual species' habitat use and habitat partitioning, and reduce potential competition for space. Cluster analysis classified trap stations into one of six habitat types that were mapped on the study area, identifying a heterogeneous assemblage of interlocking habitats. The pattern is a consequence of topographic variation on the site and, to a lesser extent, its fire history. Trapping results show P. gracilicaudatus and R. lutreolus exhibit similar macrohabitat selection, preferring topographically low habitats, with both species predominantly occupying short dense heath with dense sedge cover. The high overlap in macrohabitat use is greatly reduced when considered trap station by trap station, so that discriminant function and multiple regression analyses demonstrate marked microhabitat selection. Elevation was a highly significant variable, accounting for 41% and 27% of the variance in the habitat used by P. gracilicaudatus and R. lutreolus, respectively. This variable represents a soil moisture gradient that determines changes in the floristic and structural components of the biotic environment. Two other structural vegetation variables and vegetation height contributed 30% of the variance in P. gracilicaudatus distribution. Sedge cover was found to be significant and explained 13% of the variance in R. lutreolus distribution. Within-habitat separation was explained best with a linear combination of variables in a discriminant function, rather than by any single variable. Differential microhabitat selection, interference competition and diet separation appear to be the major factors facilitating coexistence of these two species.     

A comparative study of the seedbanks of heathland and successional habitats in Dorset, Southern England

1 Many areas of lowland heaths are being lost due to invasion by Betula spp., Pinus sylvestris, Pteridium aquilinum, Rhododendron ponticum and Ulex europaeus. One of the factors influencing the success of restoration of heathland on such sites will be the content of their viable seedbanks.
2 Ten heathland areas in the Poole Basin area of Dorset, where succession to one or more of the above species had occurred were studied. The viable seedbanks of the successional sites were compared with those of nearby heathland using Canonical Discriminant Analysis.
3 The seedbanks of all the successional stages were significantly different from the seedbank of the heath.
4 The seedbanks from the Pinus sylvestris and Pteridium aquilinum successional stages contained significantly lower numbers of heathland species than did the heathland seedbank, although few non heathland species were present.
5 The seedbanks from the Betula spp., Rhododendron ponticum and Ulex europaeus successional sites contained both significantly lower numbers of heathland species and significantly higher numbers of non heathland species than the heathland seedbank.
6 The results are discussed in relation to the restoration of heathland on successional sites and the use of the seedbank as a source of propagales for the establishment of heathland species.     

Three sympatric species of Neotoma: dietary specialization and coexistence

Summary Three sympatric species of Neotoma occur in the southern Great Basin Desert in northern Arizona. Observations and experiments from 1980–1984 focused on diet and den selection to determine to what extent woodrats partition available food and shelter. Analyses included microscopic inspection of feces from live-trapped animals, forage moisture content, and seasonal habitat utilization. Each species of woodrat was found to selectively forage on a different genus of the three evergreens on the study site: N. albigula was the only species to eat appreciable amounts of Yucca, while N. devia, specialized on Ephedra epidermis, and N. stephensi on Juniperus. Observations in the laboratory showed a linear dominance hierarchy where the larger species dominated smaller ones, i.e., N. albigula>N. stephensi>N. devia. To determine if such a hierarchy existed in the field, the behaviorally dominant species (N. albigula and N. stephensi) were continually removed (from a 25 ha experimental plot) over a 12-month period leaving only the subordinate species (N. devia) in the area. In these experiments, 40% of the dominant-species dens became occupied by 20 of the subordinate-species on the removal plot, whereas there were no interspecific den site (n=39) changes among species on the control plot. Removal of the two dominant Neotoma spp resulted in an increase of N. devia from a pre-removal high of 16 to a post-removal population of 26 individuals. These data suggest that while these woodrats may not compete for food, the subordinate species compete with the dominant species for den sites, prime dens being sequestered by the behaviorally dominant species.     

Habitat use by sympatric populations of Pseudomys novaehollandiae and Mus domesticus in coastal heathland

Abstract Patterns of habitat use by Pseudomys novaehollandiae and Mus domesticus were investigated on a 6. 8 ha area of coastal heathland, almost five years after it last burned in Myall Lakes National Park. One hundred and fifty-one trap stations were positioned on 17 traplines across a mosaic of interlocking macrohabitats. This pattern was a consequence of the site's topography and to a lesser extent fire history. Multivariate statistical procedures were employed to identify those microhabitat parameters that contribute to both individual species’ habitat use and the partitioning of habitat to reduce competition for space. Trapping results show P. novaehollandiae to be almost three times as abundant as M. domesticus at this time, although M. domesticus does show strong regional, and hence local, fluctuations in abundance over time. Both species exhibit habitat selection (P< 0. 05) with individuals preferring dry, tall closed heath and open heathland habitats. Immature P. novaehollandiae (< 15 g) were found mostly in short dense wet heath and may have been dispersing. Species exhibit high between-habitat overlap using Pianka's formula (0. 88). This overlap is substantially reduced within macrohabitats (0. 28), when trap stations are considered as resource states, suggesting a considerable degree of microhabitat separation at the trap station level. Discriminant function and multiple regression analyses identify microhabitat selection that is not strongly focused, suggesting both species to be spatial generalists. Elevation was selected as the most significant variable in these analyses and represents a soil moisture gradient that determines changes in the floristic and structural components of the biotic environment. No single habitat variable was found to explain within-habitat separation convincingly, however, when a factor analysis of habitat variables was used to produce a reduced three-dimensional factor space the outcome was marked. The 95% confidence envelopes for the distribution of each species show an extremely low overlap in factor space, with a zero overlap in the X-Y plane and less than 1% overlap in theY-Z plane. Differential micro-habitat selection does produce effective habitat partitioning, thus facilitating coexistence of these two species. We suspect this occurs primarily by M. domesticus altering habitat selection. There is evidence to suggest that M. domesticus acts as a fugitive species occupying resource niches left vacant by P. novaehollandiae and other small mammal species coexisting on the study area.     

Founder control and coexistence in a simple model of asymmetric competition for light

Size asymmetry in plant light acquisition complicates predictions of competitive outcomes in light-limited communities. We present a mathematically tractable model of asymmetric competition for light and discuss its implications for predicting outcomes of competition during establishment in two-, three-, and many-species communities. In contrast to the resource-reduction model of symmetric competition for a single resource, the model we present predicts that outcomes of asymmetric competition for light will sometimes depend on the timing of establishment and the consequent hierarchy among species in canopy position. Competitive outcomes in the model depend on the minimum light requirements (L(c)) and self-shading of species lower in the canopy compared to the light available (L(out)(*)) beneath species higher in the canopy. Succession progresses towards species with decreasing values for L(c), but arrested successions occur when initial dominants have relatively high values for L(c) but low values for L(out)(*), leading to founder control. A theoretically limitless number of species may coexist in competition for light when dominance is founder controlled. These model predictions have implications for an array of applied ecological questions, including methods to control invasive species in light-limited restored ecosystems.     

Species differences in substrate use and diet between sympatric galagos in two Kenyan coastal forests

Two sympatric species of galago,Galago zanzibaricus andG. garnettii, differed significantly in diet (judged mainly by fecal analysis) and substrate use at each of two coastal forest sites in Kenya. The larger-bodiedG. garnettii tended to use branches of a bigger diameter and to be found higher in the canopy than the smallerG. zanzibaricus. The latter species spent most of its time in the thick undergrowth. The larger species ate more fruit than the smaller. There were differences between the species in the type of invertebrate prey remains found in the feces. It is suggested that the differences in habitat use were influenced by the different body sizes of the two galagos, which so enabled these closely related species to coexist. Order of authorship is alphabetical.     

Plant coexistence in coastal heaths: Between- and within-habitat effects of competition,disturbance and predation in the post-fire environment

Abstract Previous experiments on post-fire establishment of two obligate seeders and two resprouting species suggested that initial establishment processes differ between wet- and dry-heath habitats. Disturbance, interspecific competition between seedlings and between seedlings and adults, and access for potential predators were manipulated in the field after a fire to identify mechanisms to explain between- and witm'n-habitat species coexistence. When soil surfaces are disturbed or seeds are buried, dry-heath species can establish in the wet heath. Under natural conditions, however, wet heath is rarely disturbed by animal foragers and lack of safe sites may preclude establishment of dry-heath species. In contrast, dry heath is often disturbed by ground foragers such as bandicoots and safe sites are plentiful for establishment of all heath species. Nevertheless, while wet-heath species can establish in dry heath, their seedlings are apparently unable to survive there due to the drier conditions. No evidence for competition among species was detected in either type of habitat during the first 3 years of the experiment, even at elevated seedling densities. Lack of early competition and unsaturated seed-banks may promote coexistence of species. Seedling predation from vertebrates was low and did not appear to influence patterns of coexistence within or between habitats. In terms of community theory we suggest that several interacting mechanisms are responsible for community structure in heaths after a fire. Chance and microsite disturbance are important during dispersal, while resources are critical during establishment and early survival of seedling populations. In these heath systems where fires burn extensive areas, seed-banks may be more important in temporal storage of species than patch or spatial storage as a mechanism for maintaining coexistence and plant diversity.     

Thermal benefits of artificial shelters in snakes: A radiotelemetric study of two sympatric colubrids

1.

In temperate climates, reptiles face constraining thermal conditions, and thus tradeoff predator avoidance against thermoregulatory requirements.     

Burning the Seasonal Mosaic: Preventative Burning Strategies in the Wooded Savanna of Southern Mali

Data are presented indicating a seasonal mosaic pattern of burning in the savanna of southern Mali. A seasonal mosaic is a landscape that is annually re-created by people, and which contains patches of unburned, early burned, and recently burned vegetation. A survey of over 100 farmers and in-depth interviews demonstrates that rural inhabitants of southern Mali begin an annual burning regime early in the dry season in order to fragment the landscape, with the goal of preventing later fires that can damage natural resources. The process of gradually burning off the driest vegetation creates a seasonal mosaic of habitat patches that increases the potential of the landscape for a variety of dry season land uses, including hunting, gathering of savanna products, and grazing. An analysis of a series of Landsat images shows that the practice of mosaic burning is widespread in the wooded savanna, in which burning usually begins early and large fires are rare. On the basis of recent developments in ecological theory and empirical evidence from similar burning regimes in parts of Australia, it is suggested that seasonal mosaic burning in Mali not only prevents damaging late-season fires but increases biodiversity. It is concluded that discourse on African savanna burning overemphasizes the ecologically detrimental aspects of fire, while neglecting the beneficial ones resulting in misguided policies that pose a threat to human livelihoods and savanna ecosystems.     

Seasonal succession of phytoplankton and its diversity in two highly eutrophic lakes with different conditions of stratification

Seasonal succession of phytoplankton biomass, its diversity and its photosynthetic activity in two highly eutrophic lakes have been compared. In order to test the intermediate disturbance hypothesis, the lakes have been chosen with almost the same level of trophy but different conditions of stratification, through two ice-free periods of open water with different weather conditions.High phytoplankton diversity throughout the period of investigation was characteristic for the shallower Lake Lobardzu. The number of species here was usually more than 30 and the Shannon diversity changed from 1.2 to 4.2. Owing to the frequent external disturbances, periods characterized by autogenic succession with establishing dominance and declining diversity alternated with periods of biomass reduction and rises of diversity and photosynthetic activity. In the warmer summer of 1983, with more intense warming of bottom layers and predominance of blue-greens, phytoplankton biomass was higher and diversity lower than in the cold summer of 1982.In stratified Lake Rudusku, phytoplankton diversity and number of species were usually much lower. During the long summer stratification up to three-four dominant species of blue-greens and dinoflagellates become established and competitive exclusion leading to low diversity advanced. Some changes in biomass and diversity, were caused by zooplankton activity.     

Regeneration patterns in a Central European dry heathland: effects of burning,sod-cutting and cutting

Vegetation development in dry heathlands was studied during a six-year period following experimental burning, sod-cutting and cutting in a continental area in the southern part of the Czech Republic (Podyjí National Park). Calluna vulgaris-dominated heathlands in the area were composed of uneven-aged stands and had been unmanaged for several decades. The aim of the study was testing the applicability of Western European management systems to nature conservation in a continental area. Species composition was recorded in 8 permanent plots using 25×25 cm grids, and vegetation recovery following different treatments was analysed. Calluna recovered successfully after burning. Burning dense heathlands, which contained a large amount of woody fuel, resulted in a medium-intensity fire that destroyed moss mats and litter and exposed patches of bare ground to facilitate Calluna regeneration by seed, in addition to vegetative regrowth. On the contrary, in open heathlands with patches of herbaceous vegetation, low-intensity fires which failed to expose mineral soil were more typical. Almost all Calluna regeneration was vegetative in this case, and regrowth was slower. Heathland recovery after sod-cutting to mineral soil depended on whether or not Calluna seed germination occurred in the plot. With germination, the community developed towards heathland; without towards grassland. Cutting promoted a striking increase in grass cover, which was followed by the slow recovery of Calluna. The experiments suggest that of these management systems, burning is perhaps the most appropriate in the study area. A nature conservation management system is proposed, based on sheep grazing combined with rotational burning of restricted patches.     

A possible molecular mechanism for beta-receptor desensitization: experiments and hypotheses

β-Receptor desensitization in intact NRK-S cells and in a crude membrane preparation derived from these cells was found not to involve methylation, cAMP dependent phosphorylation, Ca⁺⁺ dependent phosphorylation or ADP ribosylation, but is absolutely dependent on ATP in cell-free systems. Also, the desensitized state could not be relieved by protein phosphatases or alkaline phosphatases. Depletion of intact cells from ATP by prolonged incubation with 2-deoxy-glucose and NaN₃ did not inhibit the rapid onset of desensitization by incubating the cells with β-agonists. In order to rationalize the two seemingly contradictory findings, namely, the absolute requirement of ATP in the cell-free desensitization system and the inability to reverse the desensitized state by a variety of phosphatases, we propose that the uncoupling reaction responsible involve for establishing the desensitized state does not Our working hypothesis suggests instead, that the uncoupling reaction involves a covalent modification of the receptor protein and is catalyzed by a receptor modulator protein which is under the control of the ATP dependent phosphorylation.     

Thermal ecology of two sympatric skinks (Mabuya macrorhyncha and Mabuya agilis] in a Brazilian restinga habitat

Abstract In a restinga habitat of southeastern Brazil, we studied some aspects of the thermal biology of two sympatric skinks (Mabuya macrorhyncha and Mabuya agilis) with distinct microhabitat preferences. The two species did not differ significantly in their mean body temperature. Sex and body size did not explain body temperature variation within either species. The body temperature of both species is significantly affected by air temperature, but only that of M. agilis is significantly influenced by sand surface temperature, presumably due to its more terrestrial habit compared to M. macrorhyncha. We found no significant seasonal variation in body temperature for either species, although environmental temperatures varied seasonally, indicating that both species may be selecting an optimal range of body temperatures, possibly through behavioural thermoregulation.     

Bird communities in habitats along a successional gradient: Divergent patterns of species richness,specialization and threat

The long history of human influence on northern temperate landscapes has created a mosaic of successional stages, from closed forest to open grassland. Various species thus adapted to different habitats and it is interesting to explore how these differences in species composition among particular successional stages translate into differences at the community level. For this purpose, we surveyed breeding birds in 233 patches of five different habitats covering a gradient from bare ground to forest in 29 abandoned military training sites scattered throughout the Czech Republic. Linear mixed effects modelling revealed that late-successional habitats (dense scrubland and forest) were the most species-rich, whereas early-successional stages hosted bird communities with the highest habitat specialization and threat level. These results suggest that the habitats of late-successional stages are important for the maintenance of high bird species richness, but that early-successional habitats are essential for highly specialized and threatened bird species. Given the highly adverse impacts of agricultural intensification and land abandonment on open habitats, it is necessary to promote factors creating initial successional stages suitable for specialized and threatened species.