Spatial and reversal learning in congeneric lizards with different foraging strategies

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.     

A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications

The taxonomy of the fringe-toed lizards of the Acanthodactylus scutellatus group has long been unstable and no consensus exists on the systematic status of its various forms. A multivariate analysis of morphological characters, performed on over 1000 specimens from most of the African range of this group, allowed us to clarify the specific allocation of most of the Saharan populations included in this species group. Based on comparisons of morphology between allopatric and sympatric populations of this complex, we propose the recognition of six biological species. Our results confirm the specific status of Acanthodactylus aureus , A. dumerili , A. scutellatus, A. longipes and the recently described A. taghitensis . In addition, we re-validate A. senegalensis (occurring from Mauritania and Mali south to Senegal), which has been treated as a synonym of A. dumerili by previous authors. Acanthodactylus longipes is reported for the first time from coastal Mauritania, and A. taghitensis (previously known only from a very small region in Algeria) is reported from continental Mauritania. The systematic section of this paper includes a full list of examined material, diagnosis and known distribution of each species, in addition to some information on geographical variation and ecology. A key for specific identification is provided as an appendix.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 117−155.     

Molecular phylogeny of three Mesalina (Reptilia: Lacertidae) species (M. guttulata, M. brevirostris and M. bahaeldini) from North Africa and the Middle East: another case of paraphyly?

Mesalina is a widespread lacertid genus occurring throughout the Saharo-Sindian region from North Africa to Pakistan. It has been through a series of taxonomic revisions, but the phylogenetic relationships among the species remain unclear. In this study we estimate the phylogeographic structure of M. guttulata across most of its distributional range and we evaluate the relationships between M. guttulata and the sympatric species M. brevirostris and M. bahaeldini using partial mitochondrial DNA (mtDNA) sequences (cyt b and 16S). M. guttulata and M. brevirostris represent species complexes, whereas M. bahaeldini considered before as M. guttulata is a recently described species with very restricted distribution. Here we present the first evidence that M. guttulata is a paraphyletic taxon with respect to M. bahaeldini, while M. brevirostris proves to be a polytypic species or even a species complex, confirming previous studies. Although mtDNA markers have several properties that make them suitable for phylogeographic studies, they are not free of difficulties. Phylogeographic inferences within and between closely related species can be mislead by introgression and retention of ancestral polymorphism (incomplete lineage sorting). However, the present distribution pattern, the estimated times of divergence and the significant variation in morphology within M. guttulata led us to accept that the paraphyletic pattern observed, is most likely due to inaccurate taxonomy. Our hypothesis is that what has hitherto been considered as intraspecific variation, actually reflects species-level variation. Furthermore, our biogeographic analyses and the estimated time of divergences suggest that the present distribution of M. guttulata was the result of several dispersal and vicariant events, which are associated with historical changes (climatic oscillations and paleogeographic barriers) of late Miocene and Pliocene period.     

Influence of a broad temperature range on the oxygen consumption rates of three desert lizard species

Oxygen consumption rates of three desert lizard species: (Acanthodactylus boskianus, Lacertidae; Scincus mitranus, Scincidae; Diplometopon zarudnyi, Amphisbaenidae) were determined in relation to ambient temperatures ranging from 10 to 35 degrees C using a double-chamber, volumetric closed system. The interspecific differences in the oxygen consumption rates were found to have a close relationship between the characteristics of the M-T curves, the thermoregulatory behaviour and the ecology of the respective species. The M-T curves of the studied lizards were specifically modified and adjusted to their climatic region.     

Eating local: influences of habitat on the diet of little brown bats (Myotis lucifugus)

We employ molecular methods to profile the diet of the little brown bat, Myotis lucifugus, and describe spatial and temporal changes in diet over their maternity season. We identified 61 prey species of insects and 5 species of arachnid. The largest proportion of prey (～32%) were identified as species of the mass-emerging Ephemeroptera (mayfly) genus Caenis. Bats roosting in agricultural settings had lower dietary richness than those occupying a roost located on a forest fragment in a conservation area. We detected temporal fluctuations in diet over the maternity season. Dipteran (fly) species dominated the diet early in the season, replaced later by species of mayfly. Because our methodology provides species-level identification of prey, we were able to isolate environmental indicator species in the diet and draw conclusions about the location and type of their foraging habitat and the health of these aquatic systems. The species detected suggested that the bats use variable habitats; members of one agricultural roost foraged on insects originating in rivers or streams while those in another agricultural roost and the forest roost fed on insects from pond or lake environments. All source water for prey was of fair to good quality, though no species detected are intolerant of pollution thus the habitat cannot be classified as pristine. Our study outlines a model system to investigate the abiotic and biotic interactions between habitat factors through this simple food chain to the top predator.     

Cutting the Gordian Knot: Phylogenetic and ecological diversification of the Mesalina brevirostris species complex (Squamata,Lacertidae)

Mesalina are small lacertid lizards occurring in the Saharo‐Sindian deserts from North Africa to the east of the Iranian plateau. Earlier phylogenetic studies indicated that there are several species complexes within the genus and that thorough taxonomic revisions are needed. In this study, we aim at resolving the phylogeny and taxonomy of the M. brevirostris species complex distributed from the Middle East to the Arabian/Persian Gulf region and Pakistan. We sequenced three mitochondrial and three nuclear gene fragments, and in combination with species delimitation and species‐tree estimation, we infer a time‐calibrated phylogeny of the complex. The results of the genetic analyses support the presence of four clearly delimited species in the complex that diverged approximately between the middle Pliocene and the Pliocene/Pleistocene boundary. Species distribution models of the four species show that the areas of suitable habitat are geographically well delineated and nearly allopatric, and that most of the species have rather divergent environmental niches. Morphological characters also confirm the differences between the species, although sometimes minute. As a result of all these lines of evidence, we revise the taxonomy of the Mesalina brevirostris species complex. We designate a lectotype for Mesalina brevirostris Blanford, 1874; resurrect the available name Eremias bernoullii Schenkel, 1901 from the synonymy of M. brevirostris; elevate M. brevirostris microlepis (Angel, 1936) to species status; and describe Mesalina saudiarabica, a new species from Saudi Arabia.     

Unvarying diet of a Neotropical desert marsupial inhabiting a variable environment: the case of <Emphasis Type="Italic">Thylamys pallidior</Emphasis>

Studies on the diet and other aspects of the natural history of Neotropical marsupials have focused mainly on a few species from tropical and forest habitats. Research on desert marsupials, on the other hand, is almost absent outside Australia. The desert mouse opossum, Thylamys pallidior, is a small marsupial which inhabits the arid lands of Argentina, Chile, Bolivia, and Peru. In Argentina, it occurs in the Monte desert, a highly seasonal system with a heterogeneous landscape which affords a good opportunity to perform comparisons among different environmental situations. We analyzed fecal samples from 123 individuals and compared their diet among different seasons, habitat types, sexes, and ages. The diet of T. pallidior was composed of 68.7% of arthropods and 31.3% of plant material, mainly leaves (24.6%). We found no significant differences in the proportions of any main food category consumed for any of the factors analyzed. Consumption of both plant and animal material seems to be important in the diet of T. pallidior which manages to keep their proportions constant despite the strong variability and seasonality of the habitat. Contrary to most didelphid species, but similarly to most mammals of this area, leaves seem to represent an important source of food possibly because of their abundance and stability in the Monte desert. This represents the first study to focus on the diet of a Thylamys species inhabiting an arid environment.     

Studies of Acanthodactyla in Tunisia. II. Biometrical relations

The four species of the g. Acanthodactylus Wieg., 1834 occuring in Tunisia are quite individualized by their biometrical relations which, moreover, allow to bring out their relationships. The size of the upper quarter in our sampling oppose two groups of two species each one. The distribution of the ratio between the length of the body and that of the tail place apart A. boskianus from the three other species in a more marked way for adults of both sexes than for juveniles.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

Temporal and spatial patterns of foraging in two syntopic ant species,Manica yessensis and Formica lemani,on Mount Fuji,Japan

Spatiotemporal characteristics of terrestrial foraging were studied in two ant species, Manica yessensis and Formica lemani, in a volcanic desert on the southeast slope of Mount Fuji, Gotenba, Japan. Both ants are common in this habitat, and they construct underground nests in this dry area with sparse vegetation. Nests of M. yessensis have multiple nest-openings on the surface, whereas nests of F. lemani have very few openings, but their nesting and foraging areas overlap completely. A “mark-and-observe” method applied to M. yessensis demonstrated that worker ants of this species move between openings more than 3 m away. A study plot (6 m × 12 m quadrat) was set up, in which all nest-openings of both species were mapped. Day-long observations on numbers of foragers in this plot revealed that foraging M. yessensis are active in morning and evening, while F. lemani continues foraging all day, but both species cease activity at night. Associations between locations of foragers and nest-openings differed significantly between the two species, that is, surface foraging of M. yessensis workers was largely confined to the vicinity of their nest-openings, whereas foragers of F. lemani travelled far from their nest-openings. The function of multiple nest-openings in M. yessensis is discussed.     

Systematics and phylogeography of Acanthodactylus schreiberi and its relationships with Acanthodactylus boskianus (Reptilia: Squamata: Lacertidae)

Acanthodactylus is a widespread lacertid genus occurring from the Iberian Peninsula and western North Africa to western India including the Middle East, Cyprus, and the Arabian Peninsula. The genus is in dire need of a taxonomic revision, and the phylogenetic relationships amongst and within its species remain unclear. In particular, the taxonomy and relationship of the allopatric, narrow‐ranged Acanthodactylus schreiberi and its close relative, the widespread Acanthodactylus boskianus asper, are poorly understood. We estimated the phylogenetic and phylogeographical structure of A. schreiberi across its distribution range, and evaluated its relationships to A. b. asper, using mitochondrial and nuclear data. The phylogenetic results indicate that both species are paraphyletic, with A. schreiberi nested within A. b. asper, and the subspecies A. schreiberi syriacus nested within a distinct lineage of A. b. asper. We suggest that the group is in need of a taxonomic revision because the identified lineages and genetic diversity are incongruent with the currently recognized taxonomy. We tentatively conclude that A. schreiberi is restricted to Cyprus and Turkey, reduced to a single form, and that the populations in Lebanon and Israel belong to A. b. asper. © 2014 The Linnean Society of London     

Habitat Use and Body Mass Regulation among Warblers in the Sahel Region during the Non-Breeding Season

Migratory birds face significant challenges across their annual cycle, including occupying an appropriate non-breeding home range with sufficient foraging resources. This can affect demographic processes such as over-winter survival, migration mortality and subsequent breeding success. In the Sahel region of Africa, where millions of migratory songbirds attempt to survive the winter, some species of insectivorous warblers occupy both wetland and dry-scrubland habitats, whereas other species are wetland or dry-scrubland specialists. In this study we examine evidence for strategic regulation of body reserves and competition-driven habitat selection, by comparing invertebrate prey activity-density, warbler body size and extent of fat and pectoral muscle deposits, in each habitat type during the non-breeding season. Invertebrate activity-density was substantially higher in wetland habitats than in dry-scrubland. Eurasian reed warblers Acrocephalus scirpaceus occupying wetland habitats maintained lower body reserves than conspecifics occupying dry-scrub habitats, consistent with buffering of reserves against starvation in food-poor habitat. A similar, but smaller, difference in body reserves between wet and dry habitat was found among subalpine warblers Sylvia cantillans but not in chiffchaffs Phylloscopus collybita inhabiting dry-scrub and scrub fringing wetlands. Body reserves were relatively low among habitat specialist species; resident African reed warbler A. baeticatus and migratory sedge warbler A. schoenobaenus exclusively occupying wetland habitats, and Western olivaceous warblers Iduna opaca exclusively occupying dry habitats. These results suggest that specialists in preferred habitats and generalists occupying prey-rich habitats can reduce body reserves, whereas generalists occupying prey-poor habitats carry an increased level of body reserves as a strategic buffer against starvation.     

Foraging Behavior and Activity Patterns of Alouatta caraya in the Northeastern Argentinean Flooded Forest

Howling monkeys (Alouatta spp.) are colonizer species, showing a plasticity in behavior that allows them to inhabit different sorts of forests. There is a series of hypotheses relating demography to behavioral and ecological characteristics of howlers: (1) as howler density increases, home range size will decrease; (2) howler groups occupying small home ranges will have a high proportion of leaves in their diet; and, (3) the proportion of mature leaves in the diet is negatively correlated with group daily travel distance. To test hypotheses about howler diet and ranging pattern in relation to food resources, we studied the foraging ecology and general activity patterns of 2 groups of black howlers (Alouatta caraya) inhabiting in flooded forest of Paraná River (Argentina), the habitat with the highest density recorded for Alouatta. We found, in addition to the highest densities, also the smallest home ranges for Alouatta (Group I: 1.7 ha and Group II: 2.2 ha). However, diet was not dominated by leaves, and high quality items—flowers, fruits, new leaves—were a high proportion of the diet all year. Also, daily travel distance is not correlated with the proportion of mature leaves in the diet, but is positively correlated with the number of group confrontations. Finally, we propose a model integrating and interpreting our results as a function of the spatial and temporal distribution of high quality food resources in the Argentinean flooded forest.     

Guild mobility affects spider diversity: Links between foraging behavior and sensitivity to adjacent vegetation structure

The movement patterns of species may affect their susceptibility to modified habitat structure. It is likely that sedentary species perceive habitat features at smaller spatial extents compared to mobile species, but there is a lack of experimental research on the effects of fine-scale habitat characteristics on organisms of differing mobility. Spiders display two basic mobility levels based on foraging behavior: web-building species are restricted to specific sites whereas active hunters are mobile. We collected spiders inhabiting sagebrush shrubs with a structurally enhanced, unmodified, or removed understory, to examine (1) whether habitat structure in the immediate vicinity of shrubs affected cursorial and web spiders differently in terms of abundance and species richness and (2) which genera most contributed to changes in community composition. Shrubs without understory had reduced cursorial spider densities and species richness compared to shrubs with added and unmodified understories, whereas web spiders lacked significant responses to treatments. Community-level differences based on relative abundance of genera were detected in cursorial spiders but not in web spiders, despite a strong contribution of the web-building genus Theridion to community dissimilarities. Our results support the hypothesis that sedentary organisms may be sensitive to contiguous habitat at finer spatial scales than cursorial organisms, and highlight the risks associated with only collecting local habitat information when studying mobile species.     

Flexibility in Diet and Activity Patterns of <Emphasis Type="Italic">Macaca tonkeana</Emphasis> in Response to Anthropogenic Habitat Alteration

I examined how Sulawesi Tonkean macaques (Macaca tonkeana) respond in terms of their diet and activity patterns to anthropogenic habitat alteration in Lore Lindu National Park, Sulawesi, Indonesia. Anthropogenic habitat alteration comprised clearing of forest for agriculture and small-scale forest product collection. I quantified the diet and activity of 2 groups (CH and Anca), occupying habitats with different levels of alteration, via scan sampling. Tree abundance, key food specific density, and fruit production were greater in the minimally altered habitat (CH), substantiating the characterization of the group’s habitat as higher quality. For the group in the heavily altered habitat (Anca), alternative foods, e.g., insects, fungus, young and mature leaves, shoots, and stems, accounted for a significantly greater proportion of the diet. Dietary diversity is significantly lower in the Anca group, with 52% of their diet being palm fruits from Arenga pinnata. The activity patterns of the Anca group—more time foraging, less time moving, and more time resting than the CH group—reflect the lower resource availability in their habitat and their reliance on more alternative food items, coupled with their extremely small group size (6–9 individuals). The group may be at the optimal size in which foraging efficiency is maximized for the habitat, a response, in conjunction with dietary and behavioral flexibility, to alteration of their habitat. The results are contextualized with respect to the conservation value of human-modified landscapes.     

Northern pigtailed macaques rely on old growth plantations to offset low fruit availability in a degraded forest fragment

Space-use and foraging strategies are important facets to consider in regard to the ecology and conservation of primates. For this study, we documented movement, ranging, and foraging patterns of northern pigtailed macaques (Macaca leonina) for 14 months in a degraded habitat with old growth Acacia and Eucalyptus plantations at the Sakaerat Biosphere Reserve in northeastern Thailand. We used hidden Markov models and characteristic hull polygons to analyze these patterns in regard to fruit availability. Macaques' home range (HR) was 599 ha and spanned through a natural dry-evergreen forest (DEF), and plantation forest. Our results showed that active foraging increased with higher fruit availability in DEF. Macaques changed to a less continuous behavioral state during periods of lower fruit availability in DEF, repeatedly moving from foraging to transiting behavior, while extending their HR further into plantation forest and surrounding edge areas. Concomitantly, macaques shifted their diet from fleshy to dry fruit such as the introduced Acacia species. Our results showed that the diet and movement ecology adaptations of northern pigtailed macaques were largely dependent on availability of native fruits, and reflected a “high-cost, high-yield” foraging strategy when fresh food was scarce and dry fruit was available in plantation forest. Conversely, wild-feeding northern pigtailed macaque populations inhabiting pristine habitat approached a “low-cost, low-yield” foraging strategy. Our results outline the effects of habitat degradation on foraging strategies and show how a flexible species can cope with its nutritional requirements.     

Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

Does interspecific competition drive patterns of habitat use in desert bat communities?

Bodies of water are a key foraging habitat for insectivorous bats. Since water is a scarce and limiting resource in arid environments, bodies of open water may have a structuring effect on desert bat communities, resulting in temporal or spatial partitioning of bat activity. Using acoustic monitoring, we studied the spatial and temporal activity patterns of insectivorous bats over desert ponds, and hypothesised that sympatric bat species partition the foraging space above ponds based on interspecific competitive interactions. We used indirect measures of competition (niche overlap and competition coefficients from the regression method) and tested for differences in pond habitat selection and peak activity time over ponds. We examined the effect of changes in the activity of bat species on their potential competitors. We found that interspecific competition affects bat community structure and activity patterns. Competing species partitioned their use of ponds spatially, whereby each species was associated with different pond size and hydroperiod (the number of months a pond holds water) categories, as well as temporally, whereby their activity peaked at different hours of the night. The drying out of temporary ponds increased temporal partitioning over permanent ponds. Differences in the activity of species over ponds in response to the presence or absence of their competitors lend further support to the role of interspecific competition in structuring desert bat communities. We suggest that habitat use and night activity pattern of insectivorous bats in arid environments reflect the trade-offs between selection of preferred pond type or activity time and constraints posed by competitive interactions.     

Acanthodactyles of Tunisia: VIII--The southern acanthodactyles from south of Chott el Djerid

Applied to 135 Acanthodactylus, from eight clusters of collecting sites, a multidimensional analysis of 11 characters, mainly of colouring and scale patterns, providing 35 mathematical variables, reveals a hight intrapopulational variability. Samples dwelling on reg nebkhas and sandy knolls in the river beds are Acanthodactylus inornatus. Some specimens, in the sample from barkhanes and sand dunes areas, are morphologically close to A. longipes.     

The role of novel forest ecosystems in the conservation of wood‐inhabiting fungi in boreal broadleaved forests

The increasing human impact on the earth's biosphere is inflicting changes at all spatial scales. As well as deterioration and fragmentation of natural biological systems, these changes also led to other, unprecedented effects and emergence of novel habitats. In boreal zone, intensive forest management has negatively impacted a multitude of deadwood‐associated species. This is especially alarming given the important role wood‐inhabiting fungi have in the natural decay processes. In the boreal zone, natural broad‐leaved‐dominated, herb‐rich forests are threatened habitats which have high wood‐inhabiting fungal species richness. Fungal diversity in other broadleaved forest habitat types is poorly known. Traditional wood pastures and man‐made afforested fields are novel habitats that could potentially be important for wood‐inhabiting fungi. This study compares species richness and fungal community composition across the aforementioned habitat types, based on data collected for wood‐inhabiting fungi occupying all deadwood diameter fractions. Corticioid and polyporoid fungi were surveyed from 67 130 deadwood particles in four natural herb‐rich forests, four birch‐dominated wood pastures, and four birch‐dominated afforested field sites in central Finland. As predicted, natural herb‐rich forests were the most species‐rich habitat. However, afforested fields also had considerably higher overall species richness than wood pastures. Many rare or rarely collected species were detected in each forest type. Finally, fungal community composition showed some divergence not only among the different habitat types, but also among deadwood diameter fractions. Synthesis and applications: In order to maintain biodiversity at both local and regional scales, conserving threatened natural habitat types and managing traditional landscapes is essential. Man‐made secondary woody habitats could provide the necessary resources and serve as surrogate habitats for many broadleaved deadwood‐associated species, and thus complement the existing conservation network of natural forests.