Using scats of a generalist carnivore as a tool to monitor small mammal communities in Mediterranean habitats

Owl pellets have long been used to analyze communities of small mammals, while analogous analyses of faeces of mammal carnivores are not available. We demonstrate that common genet (Genetta genetta) scats can be used as a reliable method to sample small mammal communities and to monitor their variations. We have compiled data on 6350 small mammal remains of 18 species found in scats from 51 different latrines in a 1200 km² area of northeastern Spain. Genet scats sampled effectively 95.6% of the small mammal species ranging in size from 2.7 to 385 g. Spatial patterns of diet composition along environmental gradients of elevation, climate and land-use matched expected changes in small mammal communities along these gradients according to ecological requirements of prey species. Frequencies of occurrence of prey in genet scats were strongly correlated with frequencies of occurrence in barn owl (Tyto alba) pellets. Genet scats included two forest species not preyed upon by owls, whereas only one species was not preyed upon by genets. Forests species were more frequent in genet than in barn owl diets after correcting for environmental effects, whereas the opposite was true for open-habitat and synanthropic species. Scats of generalist carnivores can be used to estimate the spatial patterns of distribution and abundance of small mammal communities. Genet scats in fact overcome some of the limitations of more traditional sampling methods (live-trapping and owl diets), as genets were less selective and their diets reflect more accurately changes in community composition.     

The effects of landscape history and time-lags on plant invasion in Mediterranean coastal habitats

Human-driven landscape changes may promote plant invasions by increasing propagule pressure and providing favourable conditions for non-native species to establish and spread. The increase in invasion levels might not be immediate but rather exhibit a time-lag (i.e. invasion debt). Moreover, the relationship between invasion and landscape history (i.e. changes in landscape composition) might be extremely complex as it might also include extinction debts of invasive species currently in regression. In order to understand the effect of landscape history on plant invasion, we studied the invasion level in coastal plant communities affected by intense human-driven landscape alteration in the last 50 years. We identified all non-native plant species within 809 grid cells (250 × 250 m) along ~500 km of the Spanish SW coast. We tested the association of (1) non-native richness conditional on the overall presence of non-natives (at least one species), and (2) the occurrence of the most frequent non-native species, with the percentage of human land-cover categories in 1956, 1991 and 2007 using a multimodel inference approach. We used these models to project future invasion patterns in the region. We found non-native richness to be more associated with land-cover variables in 1956 than in 2007, suggesting the existence of an invasion debt, but not for the overall presence of non-natives. For most frequent species, the effects of past landscape alteration depended on the introduction pathway and the region of origin. Species used in agriculture were more related to past cropland area, while most of the species used in gardening or forestry showed higher affinity for recently altered areas. These results highlight the relevance of the species’ introduction history and landscape history in assessing future long-term invasion trends.     

Sexual segregation of seasonal foraging habitats in a non-migratory marine mammal

Many animal species segregate by sex. Such segregation may be social in nature, or ecological, or both. Grey seals (Halichoerus grypus), like many large mammals, are sexually size dimorphic. In size dimorphic species, allometric differences in morphology, metabolic rate and reproductive costs are likely. Such differences may require the sexes to use different foraging strategies or different habitats. To investigate sexual segregation of habitat in grey seals, we used satellite tracks from 95 (male 46; female 49) adults breeding at Sable Island, Nova Scotia (44 degrees N, 60 degrees W) collected from 1995 to 2005. Location estimates were made from satellite fixes using a state-space movement model to estimate true locations and regularize them in time. Location estimates were used to calculate home range kernels of male and female habitat use each month. Month by sex kernel home ranges revealed striking differences and dynamics in habitat use between males and females on spatial scales broader than most terrestrial examples and at temporal and spatial resolutions rarely available for marine species. Differences were most pronounced just before (October-December) and immediately after breeding (February-March). During both periods, males primarily used areas along the continental shelf break, while females mainly used mid-shelf regions. Coupled with previously identified sex-specific seasonal patterns of energy storage, diving and diet, our findings suggest that males and females differ profoundly in their spatial foraging strategies. These differences may serve to maximize fitness by reducing intersexual competition during key foraging periods.     

Small mammal community ecology

Summary The abundance of food in natural 1-ha plot of shortgrass prairie was artificially manipulated by adding alfalfa pellets and whole oats on a regular schedule. The small mammal species naturally inhabiting the manipulated food plot did not respond in either their density or weight to the supplemented food. However, a new specialized seed-eating species, Dipodomys ordii, invaded the food plot and persisted in relatively high density. As a result of this colonization, species diversity was significantly higher on the food plot relative to the unmanipulated control plot. In a second experiment, application of water and nitrogen to two 1-ha plots of shortgrass prairie resulted in increased productivity. However, the increased production on this latter treatment was associated with vegetation growth and thus major changes in habitat structural characteristics relative to the control treatment. Two new species, Microtus ochrogaster and Reithrodontomys megalotis, colonized the nitrogen+water treatment, but other small mammal species resident to the shortgrass prairie largely avoided this treatment. As a result of this manipulation, species diversity was significantly lower than the species diversity on the control treatment. These two results agree with the MacArthur prediction regarding how species diversity responds to an increase in production of scarce resources without changes in other habitat variables, and to an increase in productivity associated with changes in other habitat variables.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Multi-scale analysis of butterfly diversity in a Mediterranean mountain landscape: mapping and evaluation of community vulnerability

This paper is an attempt to outline a protocol for animal diversity census and evaluation aimed for areas in view of landscape planning of territories of hundred square kilometres and more, that may work utilising different faunal groups and be anyway useful at various scales. Many papers are addressed to elaborate tools for landscape planning starting from biodiversity evaluation and butterflies are often utilised because of their sensitivity to landscape modifications. In this work, the biodiversity evaluation has been performed using three hierarchically linked landscape units at micro-, meso- and macroscale. Being species diversity values often inadequate to define the conservation interest of a landscape portion, more importance has been given to which species compose the species assemblages. A community vulnerability Index was coded and used for evaluating potential consequences of human disturbance on butterfly assemblages. Forty-four year samples were gained by visual census in the Sila Greca, Southern Italy, on an area of approximately 520 square kilometres. During 5 years work, 2,535 specimens and 94 species were recorded, equal to 75.8% of the whole Calabrian fauna. Four vulnerability levels have been established and used for mapping butterfly assemblage vulnerability in the area, starting from a vegetation map. Species richness was found somewhat contradictory at micro-scale, where the community vulnerability Index gives a sounder approach. S diversity gives a more reliable picture of naturalness at meso-scale, a level we identified with the “ecotope”. At this more “geomorphic” scale level, biological functions reflected by butterfly assemblages revealed to be clearly linked to seral processes. Similarity analysis results show that the ecotope species richness, here called “eta-diversity”, could be an useful measure of zoological landscape (faunation) potentialities.     

Identification of vertebrate scavengers of small mammal carcasses in a forested landscape

We identified vertebrate scavengers of small mammal carcasses at the 780-km² Savannah River Site during the winter of 2000–2001. Rodent carcasses, differing in size and visual conspicuousness, were placed in upland pine forests and bottomland hardwood forests during six 2-week periods. Sixty-two of the 96 carcasses (65%) were removed by vertebrates. With the aid of remote photography, we identified 11 species of scavengers removing carcasses. RaccoonsProcyon lotor, gray foxesUrocyon cinereoargenteus, and feral pigsSus scrofa scavenged most frequently. The mean elapsed time for carcass removal was 5.6 days. The number of carcasses removed by vertebrates did not differ significantly with respect to carcass size, visual conspicuousness, or habitat type; however, air temperature was strongly correlated (positively) with carcass removal. Our study demonstrates that many mammal species are capable of utilizing small carrion items as a food resource, and suggests that scavenging may account for a higher proportion of the diet of some facultative scavengers than is now widely assumed.     

Contrasting life histories in neighbouring populations of a large mammal

Background

A fundamental life history question is how individuals should allocate resources to reproduction optimally over time (reproductive allocation). The reproductive restraint hypothesis predicts that reproductive effort (RE; the allocation of resources to current reproduction) should peak at prime-age, whilst the terminal investment hypothesis predicts that individuals should continue to invest more resources in reproduction throughout life, owing to an ever-decreasing residual reproductive value. There is evidence supporting both hypotheses in the scientific literature.

Methodology/Principal Findings

We used an uncommonly large, 38 year dataset on Alpine chamois (Rupicapra rupicapra) shot at various times during the rutting period to test these two hypotheses. We assumed that body mass loss in rutting males was strongly related to RE and, using a process-based approach, modelled how male relative mass loss rates varied with age. For different regions of our study area, we provide evidence consistent with different hypotheses for reproductive allocation. In sites where RE declined in older age, this appears to be strongly linked to declining body condition in old males. In this species, terminal investment may only occur in areas with lower rates of body mass senescence.

Conclusions/Significance

Our results show that patterns of reproductive allocation may be more plastic than previously thought. It appears that there is a continuum from downturns in RE at old age to terminal investment that can be manifest, even across adjacent populations. Our work identifies uncertainty in the relationship between reproductive restraint and a lack of competitive ability in older life (driven by body mass senescence); both could explain a decline in RE in old age and may be hard to disentangle in empirical data. We discuss a number of environmental and anthropogenic factors which could influence reproductive life histories, underlining that life history patterns should not be generalised across different populations.     

Invasion of alien macroalgae in different Mediterranean habitats

The study evaluated different macroalgal invasions in the main Mediterranean coastal habitats on hard bottom. Biodiversity, species composition and structure of macroalgal assemblages were compared among non-invaded areas and areas invaded by the Chlorophyta Caulerpa racemosa var. cylindracea and by the turf-forming Rhodophyta Womersleyella setacea in three different habitats: shallow rocky bottom, deep rocky bottom and dead matte of the seagrass Posidonia oceanica. Results showed that alien macroalgae constituted a relevant component of benthic assemblages in invaded areas of the Mediterranean Sea. Assemblages invaded by Womersleyella setacea and Caulerpa racemosa showed lower values of diversity and large differences in the structure and species composition related to non-nvaded assemblages. The species that mostly suffered from invasion were erect species reproducing sexually; moreover, the dominance of W. setacea led to low abundance of native filamentous algae, while C. racemosa colonization seemed particularly threatening for encrusting algae. All the studied habitats appeared highly invasible by alien macroalgae, even if W. setacea appeared more invasive in deeper habitats, while colonization of C. racemosa seemed more serious in shallower habitats; the dead matte of P. oceanica represented a suitable substrate for the spread of both species. Differences among assemblages in different habitats were reduced in invaded areas.     

Butterfly community structure in fragmented habitats

We analysed effects of habitat fragmentation on the diversity, abundance, and life history traits of butterflies on 33 calcareous grasslands. Diversity of butterflies was positively correlated with habitat area (as was plant diversity), but not with habitat isolation. In contrast to expectations, butterfly densities of polyphagous and oligophagous species declined with habitat area whereas densities of monophagous species increased. The z -values, i.e. the slope of species–area relationships, increased with food plant specialization, from 0.07 in polyphagous, 0.11 in oligophagous, 0.16 in strongly oligophagous to 0.22 in monophagous species, and were 0.14 in plant species. Significant z -values were not only found for total species richness, based on a sample size adjusted to fragment area ( z  = 0.12), but also for the local density of butterfly species richness, based on equal sample size across all habitat fragments ( z  = 0.06). To our knowledge, this is the first study to show differential responses of monophagous, oligophagous and polyphagous species to area with respect to species richness and population density.     

A long term study of small mammal populations in a Brazilian agricultural landscape

Long-term monitoring of small mammal populations is very important to understand the variations in temporal abundance on a large time scale, which are related to ecological, economic and epidemiological phenomena. The aim of this study is to monitor the populations of the marsupials Didelphis aurita and Philander frenatus and the rodents Nectomys squamipes, Akodon cursor and Oligorysomys nigripes in a locality of typical Brazilian rural landscape, Sumidouro Municipality, Rio de Janeiro State, Brazil. A mark-recapture study was conducted during five years. We analyzed the population dynamics, the reproduction and age structure of these species. Both marsupials presented higher population sizes in the end of wet period and beginning of the dry period, which can be explained by the seasonal reproduction which begins in the middle of the dry period and ends in the last months of the wet period. N. squamipes reproduced throughout the year but mostly during rainy periods, due to the close association of this rodent to resources found in the water. Higher survivorship and recruitment rates were in the end of the wet season. The rodent A. cursor had an opportunist reproduction, resulting in high turnover rates. Survivorship increased with the effects of the dry periods. O. nigripes showed a clear annual pattern of population cycle with peaks during the dry season. The rodents did not show potential to present outbreaks and become agricultural pests. The annual population cycles of O. nigripes and the unique peak of A. cursor population during five years highlight attention to their importance as wild reservoirs of the hantavirus disease. Their ecological characteristics associated to their opportunistic behavior make these species prone to be good reservoirs of zoonoses.     

Towards a standardized index of European rabbit abundance in Iberian Mediterranean habitats

European rabbits Oryctolagus cuniculus are a keystone species in Iberian Mediterranean ecosystems. However, the reliability of methods for estimating rabbit abundance, particularly when at low numbers, is not well understood. Further, better standardization of these methodologies would allow abundance estimates to be more reliably compared between areas and periods. Consequently, we compared several frequently used methods of estimating rabbit abundance and assessed their advantages and disadvantages. During the summers of 2008 and 2009, in 11 localities of central-southern Spain we undertook (a) driving transect counts of rabbits, either at dusk or at night, (b) linear transects on foot recording rabbit signs, (c) cleared-plot pellet counts at permanent plots, and (d) standing crop counts, both with and without habitat stratification. Density estimated at night from driving transects using the Distance Sampling method (the reference method against which all other indices were compared) varied from 0 to 2.69 rabbits ha⁻¹. Most pellet-count indices were significantly related to the reference method. In particular, cleared-plot pellet counts in permanent plots corrected for pellet persistence showed the best correlation with the reference method. In contrast, latrine counts were not related to the reference method index, and we recommend against their use. A standard methodology based on cleared-plot pellets counts could be used to monitor rabbit abundance on a large scale.     

Temporal shifts as elusive responses to anthropogenic stressors in a mammal community

Conservation decision-making often relies on species’ distribution changes in response to anthropogenic disturbances but overlook their temporal responses. Filling the knowledge gap on the temporal shifts as elusive responses of wild animals to human activity is important because this may provide information for more proactive conservation planning. In this study, we used camera traps in a field survey technique to investigate the trade-offs between spatial and temporal responses of a mammal community to major human activities in Qinling Mountains, China. We focused on five most abundant mammalian species including giant panda (Ailuropoda melanoleuca), takin (Budorcas taxicolor), wild boar (Sus scrofa), tufted deer (Elaphodus cephalophus), and Chinese goral (Naemorhedus griseus), and examined the drivers of, and the trade-offs between the change of their spatial distributions and daily activity patterns in response to anthropogenic disturbances. We found that in response to human interferences, giant panda and takin mainly changed their distributions, while wild boar, tufted deer and Chinese goral altered their daily activity patterns, indicating the elusive responses of the latter species under anthropogenic stressors. In addition, anthropogenic stressors such as farming and tourism have more profound impacts on mammal communities than previously revealed by species distribution modeling only. For nature reserves that aim to conserve multiple species simultaneously, a more flexible, adaptive management framework is thus needed to capture the trade-offs between multiple species’ spatial and temporal responses to anthropogenic disturbance.

     

Phylogenetic structure of mammal assemblages at large geographical scales: linking phylogenetic community ecology with macroecology

Phylogenetic community ecology seeks to explain the processes involved in the formation of species assemblages by analysing their phylogenetic structure, and to date has focused primarily on local-scale communities. Macroecology, on the other hand, is concerned with the structure of assemblages at large geographical scales, but has remained largely non-phylogenetic. Analysing the phylogenetic structure of large-scale assemblages provides a link between these two research programmes. In this paper, I ask whether we should expect large-scale assemblages to show significant phylogenetic structure, by outlining some of the ecological and macroevolutionary processes that may play a role in assemblage formation. As a case study, I then explore the phylogenetic structure of carnivore assemblages within the terrestrial ecoregions of Africa. Many assemblages at these scales are indeed phylogenetically non-random (either clustered or overdispersed). One interpretation of the observed patterns of phylogenetic structure is that many clades underwent rapid biome-filling radiations, followed by diversification slowdown and competitive sorting as niche space became saturated.     

Avian community composition of kopjes in a heterogeneous landscape

We examined avian communities of kopjes (naturally occurring insular rock outcrops) in grassland and thorn tree woodland habitats in the Seronera region of Serengeti National Park, northwestern Tanzania. Although kopjes cover a small proportion of the Park's area, they provide resources that are uncommon in the Serengeti landscape and are known to host diverse, yet poorly documented, biotic communities. The primary objectives of this study were (1) to compare avian communities of kopjes with those of their surrounding habitat matrixes; (2) to compare the avian communities among kopjes; and (3) to determine the effects of kopje characteristics (e.g., size, vegetation cover, level of human disturbance and matrix type) on avian diversity and community composition. The avian communities of kopjes differed significantly from those of the matrixes in species composition and guild characteristics. Rare species, frugivorous species and nectarivorous species were more abundant on kopjes, whereas there were more ground-feeding species in matrix sites. Species richness was positively correlated with the area of kopjes covered by tall vegetation (5+ m), but neither total habitat area nor total vegetation cover significantly affected avian diversity. The surrounding habitat type, the fruiting phenology of Ficus and Commiphora trees and the level of human disturbance also influenced the presence and abundance of individual species and accounted for differences in community composition among kopjes. Our results show that kopjes support unique avian assemblages comprising many species that are otherwise rare in Serengeti, and consequently that kopjes may be local hotspots of avian diversity in the region.     

Degree of landscape fragmentation influences genetic isolation among populations of a gliding mammal

Forests and woodlands are under continuing pressure from urban and agricultural development. Tree-dependent mammals that rarely venture to the ground are likely to be highly sensitive to forest fragmentation. The Australian squirrel glider (Petaurus norfolcensis) provides an excellent case study to examine genetic (functional) connectivity among populations. It has an extensive range that occurs in a wide band along the east coast. However, its forest and woodland habitat has become greatly reduced in area and is severely fragmented within the southern inland part of the species' range, where it is recognised as threatened. Within central and northern coastal regions, habitat is much more intact and we thus hypothesise that genetic connectivity will be greater in this region than in the south. To test this we employed microsatellite analysis in a molecular population biology approach. Most sampling locations in the highly modified south showed signatures of genetic isolation. In contrast, a high level of genetic connectivity was inferred among most sampled populations in the more intact habitat of the coastal region, with samples collected 1400 km apart having similar genetic cluster membership. Nonetheless, some coastal populations associated with urbanisation and agriculture are genetically isolated, suggesting the historic pattern observed in the south is emerging on the coast. Our study demonstrates that massive landscape changes following European settlement have had substantial impacts on levels of connectivity among squirrel glider populations, as predicted on the basis of the species' ecology. This suggests that landscape planning and management in the south should be focused on restoring habitat connectivity where feasible, while along the coast, existing habitat connectivity must be maintained and recent losses restored. Molecular population biology approaches provide a ready means for identifying fragmentation effects on a species at multiple scales. Such studies are required to examine the generality of our findings for other tree-dependent species.     

Habitat structure and disturbance affect small mammal populations in Mediterranean forests

Habitat structure, the physical arrangement of objects in space, affects animal populations, ecological interactions, species diversity, and ecosystem functioning. Conventional forest management practices that reduce habitat structural complexity are thus under reappraisal, and there are increasing efforts to understand how habitat structure affects functionally important species. We investigate here the effect of habitat structures on small mammal dens and their interactions with widespread antagonistic species (wild boars). Among habitat structures we considered tree dimension (diameter), shrub cover, rock cover, dead wood volume, and stump area. Ground cover with rooting signs measured wild boar disturbance. The number of small mammal dens was related positively to rock cover and tree diameter, and negatively to wild boar disturbance. Additionally, a positive interaction between wild boar disturbance and tree diameter emerged, showing that the effect of big trees on small mammal nesting was more evident with higher levels of disturbance. This study suggests that habitat structures that can be affected by forest management provide effective refuges for functionally important species. The effect of habitat structure on species survival and interactions thus needs to be addressed to understand ecosystem management and functioning.