The Neotropical otter Lontra longicaudis: a comprehensive update on the current knowledge and conservation status of this semiaquatic carnivore

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The macroevolutionary relationship between diet and body mass across mammals

Body mass and diet are two fundamental ecological parameters that influence many other aspects of an animal's biology. Thus, the potential physiological and ecological processes linking size and diet have been the subject of extensive research, although the broad macroevolutionary relationship between the two traits remains largely unexplored phylogenetically. Using generalized Ornstein–Uhlenbeck models and data on over 1350 species of mammal, we reveal that evolutionary changes in body mass are consistently associated with dietary changes across mammals. Best‐fitting models find that herbivores are substantially heavier than other dietary groups and that omnivores are frequently intermediate in mass between herbivores and carnivores. Interestingly, although flying and swimming both place very different physical constraints on mass, bats still follow the general mammalian pattern but marine mammals do not. Such differences may be explained by reduced gravitational constraints on size in water along with ecological differences in food availability between aquatic and terrestrial realms, allowing marine carnivores to become the largest mammals on earth. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 173–184.     

Body size of insular carnivores: evidence from the fossil record

Aim Our goals here are to: (1) assess the generality of one aspect of the island rule – the progressive trend towards decrease in size in larger species – for fossil carnivores on islands; (2) offer causal explanations for this pattern and deviations from it – as far as fossil carnivores are concerned; and (3) estimate the speed of this trend. Location Oceanic and oceanic‐like islands world‐wide. Methods Body size estimates of fossil insular carnivores and of their phylogenetically closest mainland relative were obtained from our own data and the published literature. Our dataset consisted of 18 species from nine islands world‐wide. These data were used to test whether the body size of fossil insular carnivores varies as a function of body size of the mainland species in combination with characteristics of the island ecosystem. Results Dwarfism was observed in two canid species. Moderate decrease in body mass was observed in one hyena species. Gigantism was observed in one otter species. Moderate body mass increase was observed in two otter species, one galictine mustelid and perhaps one canid. Negligible or no change in body mass at all was observed in five otter species, three galictine mustelids and one genet. Size changes in teeth do not lag behind in comparison to skeletal elements in the dwarfed canids. The evolutionary speed of dwarfism in a canid lineage is low. Main conclusions Size change in fossil terrestrial insular carnivores was constrained by certain ecological conditions, especially the availability of prey of appropriate body size. When such alternative prey was not available, the carnivores retained their mainland size. The impact of competitive carnivores seems negligible. The case of (semi‐)aquatic carnivores is much less clear. The species that maintained their ancestral body mass may have changed their diet, as is evidenced by their dentition. Among the otters, one case of significant size increase was observed, perhaps best explained as being due to it entering the niche of an obligate aquatic otter. Dwarfism was not observed in otters. The island rule seems to apply to fossil carnivores, but with exceptions. The dependency of the island rule on resource availability is emphasized by the present study.     

Reproductive system,social organization,human disturbance and ecological dominance in native populations of the little fire ant,Wasmannia auropunctata

The invasive ant species Wasmannia auropunctata displays both ecologically dominant and non‐dominant populations within its native range. Three factors could theoretically explain the ecological dominance of some native populations of W. auropunctata: (i) its clonal reproductive system, through demographic and/or adaptive advantages; (ii) its unicolonial social organization, through lower intraspecific and efficient interspecific competition; (iii) the human disturbance of its native range, through the modification of biotic and abiotic environmental conditions. We used microsatellite markers and behavioural tests to uncover the reproductive modes and social organization of dominant and non‐dominant native populations in natural and human‐modified habitats. Microsatellite and mtDNA data indicated that dominant and non‐dominant native populations (supercolonies as determined by aggression tests) of W. auropunctata did not belong to different evolutionary units. We found that the reproductive system and the social organization are neither necessary nor sufficient to explain W. auropunctata ecological dominance. Dominance rather seems to be set off by unknown ecological factors altered by human activities, as all dominant populations were recorded in human‐modified habitats. The clonal reproductive system found in some populations of W. auropunctata may however indirectly contribute to its ecological dominance by allowing the species to expand its environmental niche, through the fixation over time of specific combinations of divergent male and female genotypes. Unicoloniality may rather promote the range expansion of already dominant populations than actually trigger ecological dominance. The W. auropunctata model illustrates the strong impact of human disturbance on species’ ecological features and the adaptive potential of clonal reproductive systems.     

Brazilian populations of Drosophila maculifrons (Diptera: Drosophilidae): low diversity levels and signals of a population expansion after the Last Glacial Maximum

The Quaternary period was marked by considerable changes in climate. Such palaeoclimatic changes affected the population dynamics of many species, both in the Northern and in the Southern Hemisphere. However, the extent of these impacts on the demographic patterns of Neotropical species presenting different ecological requirements remains unclear. Drosophila maculifrons DUDA 1947 belongs to the guaramunu group of Drosophila and represents a potential indicator of the genetic consequences caused by the climatic fluctuations of the Quaternary, because it seems to be sensitive to temperature and humidity shifts. The aim of this study was to evaluate the evolutionary processes subjacent to the patterns of intraspecific diversity and structure of different populations of D. maculifrons. In total, 152 individuals were collected in the south and south‐east Brazil. Phylogenetic and phylogeographical analyses were performed based on sequences of COI and COII mitochondrial genes. In general, the results pointed to Brazilian populations of D. maculifrons being extremely impoverished in terms of mitochondrial diversity and population structure, which could be explained by a recent population expansion event dated to approximately 12 000 years ago. In fact, with the assistance of species palaeo‐distribution modelling strategies, it was possible to infer that most of the sampled region did not present the D. maculifrons environmental suitability requirements at least during the period of the Last Glacial Maximum. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 55–66.     

Niche dynamics in the European ranges of two African carnivores reflect their dispersal and demographic histories

According to recent phylogeographical evidence, the common genet (Genetta genetta) and the Egyptian mongoose (Herpestes ichneumon) have dissimilar dispersal histories from Maghreb to south‐western Europe. Through comparative ecological niche modelling based on >1100 occurrences, we assessed whether the niche dynamics (i.e. niche shift versus conservatism) of the two species in their European ranges reflected DNA‐based demographic scenarios. Sensitivity analyses and projections of climatic niche models from the species' native ranges (Africa and Middle East) to Europe yielded support for (1) partial climatic niche shift in the northern European range of the common genet and (2) climatic niche conservatism in the Egyptian mongoose. Our results were consistent with demographic scenarios that predicted multiple introductions and demographic expansion in the common genet and long‐term, stable historical demography in the Egyptian mongoose. Our models further predicted a range expansion of the common genet in north‐western France and Italy, and progression of the Egyptian mongoose into Europe from the Near East. Overall, our study suggested a scenario of different niche dynamics in Europe for these two species of African carnivores, supporting the view that historical factors such as dispersal and demographic history may shape niche dynamics and thus distribution potential in colonized ranges. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 737–751.     

A link between historical population decline in the threatened great bustard and human expansion in Iberia: evidence from genetic and demographic data

Early anthropogenic impacts on the abundance and distribution of wild species are difficult to document, but can help us to understand the causes and relative importance of current declines. Genetic data can be of use in inferring historical demographic events, but the accuracy of these inferences depends on the availability and precision of demographic parameters that are difficult to obtain in the field. Here, we use demographic data on Iberian populations of the threatened great bustard (Aves: Otis tarda), obtained from an intensive population monitoring programme over the last 20 years, to estimate critical population parameters (population size and generation time), which are then used in a Bayesian Skyline Plot (BSP) analysis of mitochondrial DNA sequence data to assess changes in population size over the last several thousand years. BSP showed a sudden and sharp great bustard population decline coinciding with human expansion in Iberia, and the associated agricultural and urban development and increased hunting pressure. These results illustrate the importance of human population size as a possible ultimate cause of an environmental impact that occurred in the historical past, a fact that has often been neglected. Our results also suggest the role of human activities in driving historical population declines in great bustards, and underscore the importance of precise, long‐term field data to infer past demographic trends from parameters of extant populations. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 518–527.     

Latitude drives diversification in Madagascar's endemic dry forest rodent Eliurus myoxinus (subfamily Nesomyinae)

Numerous hypotheses have been proposed for the historical processes governing the rich endemism of Madagascar's biodiversity. The ‘watershed model’ suggests that drier climates in the recent geological past have resulted in the contraction of forests around major watersheds, thereby defining areas of endemism. We test whether this hypothesis explains phylogeographical patterns in a dry forest‐dependent rodent, Eliurus myoxinus, an endemic species widely distributed through western Madagascar. We sequenced the mitochondrial cytochrome b locus and nuclear introns of the β‐fibrinogen and the growth hormone receptor genes for E. myoxinus. Using a parametric bootstrapping approach, we tested whether the mitochondrial gene tree data fit expectations of local differentiation given the watershed model. We additionally estimated population differentiation and historical demographic parameters, and reconstructed the spatial history of E. myoxinus to highlight spatial and temporal patterns of differentiation. The data do not support the watershed model as a clear explanation for the genetic patterns of diversity within extant E. myoxinus populations. We find striking patterns of latitudinal genetic structure within western Madagascar, and indicate possible roles for environmental and ecological gradients along this axis in generating phylogeographical diversity. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 500–517.     

Do non‐native fish as prey favour the conservation of the threatened indigenous Eurasian otter?

1. Biological invasions are considered a major threat to biodiversity. Most research has focused on the distribution, biology and impacts of non‐native species on native fauna and flora. However, few studies have explored their role as prey for native predators of conservation concern. 2. To assess the incidence and intensity of predation by the Eurasian otter Lutra lutra on established non‐native fish species, data were collated from the published literature. To be selected, studies had to cover at least 1 year, analyse more than 100 spraints and report the study period and percentage relative frequency (%RF) of all prey fish species. 3. To permit reliable, time‐related comparisons with %RF of non‐native fishes in otter diet, we also reviewed available information about both the distribution of non‐native fishes and history of their introductions to European countries, revealing a decrease with longitude in the number of naturalised non‐native fishes taken (ranging between 5 and 34) and their percentage in each fish assemblage. 4. Our selective criteria were met by 30 dietary studies from 44 study areas in 15 European countries during 1970–2010. The extent to which otters rely on non‐native fishes was almost negligible (mean %RF = 4.8), with the number of non‐native fishes preyed upon by otters decreasing with both latitude and longitude. 5. The %RF of non‐native fish in the diet increased slightly with time, with otters preying significantly more on non‐native fish in study areas where alterations of the fish assemblage had been highlighted in the reference papers. No relationship was found between otter diet breadth and the occurrence of non‐native fishes in their diet. 6. The current role of non‐native species in otter diet suggests that effective otter conservation management plans should focus on the maintenance and/or enhancement of native fish assemblages.     

Phylogeography of the Trans‐Volcanic bunchgrass lizard (Sceloporus bicanthalis) across the highlands of south‐eastern Mexico

We quantify the population divergence processes that shaped population genetic structure in the Trans‐Volcanic bunchgrass lizard (Sceloporus bicanthalis) across the highlands of south‐eastern Mexico. Multilocus genetic data from nine nuclear loci and mitochondrial (mt)DNA were used to estimate the population divergence history for 47 samples of S. bicanthalis. Bayesian clustering methods partitioned S. bicanthalis into three populations: (1) a southern population in Oaxaca and southern Puebla; (2) a population in western Puebla; and (3) a northern population with a broad distribution across Hidalgo, Puebla, and Veracruz. The multilocus nuclear data and mtDNA both supported a Late Pleistocene increase in effective population size, and the nuclear data revealed low levels of unidirectional gene flow from the widespread northern population into the southern and western populations. Populations of S. bicanthalis experienced different demographic histories during the Pleistocene, and phylogeographical patterns were similar to those observed in many co‐distributed highland taxa. Although we recommend continuing to recognize S. bicanthalis as a single species, future research on the evolution of viviparity could gain novel insights by contrasting physiological and genomic patterns among the different populations located across the highlands of south‐eastern Mexico. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 852–865.     

Dietary complexity and hidden costs of prey switching in a generalist top predator

1. Variation in predator diet is a critical aspect of food web stability, health, and population dynamics of predator/ prey communities. Quantifying diet, particularly among cryptic species, is extremely challenging, however, and differentiation between demographic subsets of populations is often overlooked.
2. We used prey remains and data taken postmortem from otter Lutra lutra to determine the extent to which dietary variation in a top predator was associated with biotic, spatial, and temporal factors.
3. Biotic data (e.g., sex, weight, and length) and stomach contents were taken from 610 otters found dead across England and Wales between 1994 and 2010. Prey remains were identified to species where possible, using published keys and reference materials. Multi‐model inference followed by model prediction was applied to test for and visualize the nature of associations.
4. Evidence for widespread decline in the consumption of eels (Anguilla anguilla) reflected known eel population declines. An association between eel consumption and otter body condition suggested negative consequences for otter nutrition. Consumption of Cottus gobio and stickleback spp. increased, but was unlikely to compensate (there was no association with body condition). More otters with empty stomachs were found over time. Otter sex, body length, and age‐class were important biotic predictors of the prey species found, and season, region, and distance from the coast were important abiotic predictors.
5. Our study is unique in its multivariate nature, broad spatial scale, and long‐term dataset. Inclusion of biotic data allowed us to reveal important differences in costs and benefits of different prey types, and differences between demographic subsets of the population, overlaid on spatial and temporal variation. Such complexities in otter diet are likely to be paralleled in other predators, and detailed characterization of diet should not be overlooked in efforts to conserve wild populations.

     

The Scotch broom,Cytisus scoparius (Fabaceae), a paradox in Denmark – an invasive plant or endangered native species?

Scotch broom, Cytisus scoparius, spreads rapidly in parts of Denmark and is considered an invasive species by some authors. However, the species has been present in the Danish flora for centuries and is therefore considered native to Denmark. In the present study we explore whether Danish Scotch broom consists of one or two gene pools with potential differences in phenotype and invasiveness. One plastid and five nuclear microsatellite markers were used to reveal potential substructuring of Danish Scotch broom. Nine populations were included representing populations exhibiting invasive behaviour and populations showing non‐invasive behaviour. An Italian population was used as reference. Bayesian analysis based on genetic markers indicated that the sampled populations form two distinct gene pools, and this pattern was supported by neighbour‐joining trees. Measurements of height and width of the analysed plants showed that the two gene pools correspond to populations exhibiting either a horizontal habit and non‐invasive behaviour or an erect habit and, in some cases, invasive behaviour. The Italian population clustered with the erect ones. We discuss the origin and taxonomic status of the two gene pools and conclude that Danish horizontal Scotch broom should be given a formal taxonomic status in order to initiate conservation activities for its protection. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Non‐native conditions favor non‐native populations of invasive plant: demographic consequences of seed size variation?

Trait differences between native and non‐native populations may explain the greater abundance and impact of some organisms in their non‐native ranges than in their native ranges. Here, we conducted reciprocal common gardens in southwestern Turkey (home) and central Argentina (away) to explore the hypothesis that the greater success of the invasive ruderal Centaurea solstitialis in Argentina than Turkey is partially explained by differences between home and away populations. Unusual among common gardens, our experimental design included seed additions to explicitly evaluate population level responses, as well as disturbance and no‐disturbance treatments. We documented seed mass in native and non‐native populations, and during the experiment, we periodically measured density, plant size, and herbivory. After six months, we determined the establishment of plants for populations from both origins in both home and away common gardens. Seed mass was two times larger for Argentinean than Turkish populations. Density, plant size and final establishment were also greater for plants from Argentinean than from Turkish populations, but only in the common garden in Argentina. In Turkey, no differences between population origins were detected for these variables. Herbivory was similar for populations from both origins in both common gardens. As expected, disturbance generally increased plant performance in both regions. Our results suggest that increased seed size in non‐native populations may have demographic consequences under non‐native conditions that can contribute to the invasive success of C. solstitialis. This is the first reciprocal common garden that supports the idea that seed size variation contributes to demographic differences for an invasive species between native and non‐native distributions, but our findings further suggest that seed size effects on demography depend on the ecological context in which population processes occur.     

The effects of the Late Quaternary glacial–interglacial cycles on Anatolian ground squirrels: range expansion during the glacial periods?

The present study aimed to understand how Anatolian ground squirrels, Spermophilus xanthoprymnus (Bennett, 1835), have responded to global climate changes through the Late Quaternary glacial–interglacial cycles. Accordingly, ecological niche modelling was used, together with molecular phylogeography. Using species occurrence data compiled from field observations and relevant sources and the maximum entropy machine learning algorithm in MAXENT, an ecological niche model was developed to predict the potential geographical distribution of S. xanthoprymnus under reconstructed past (the Last Interglacial, approximately 130 000–116 000 years ago and the Last Glacial Maximum, 21 000 years ago) and present (1950–2000) bioclimatic conditions. In addition, using cytochrome b mitochondrial DNA sequences deposited in GenBank and the Bayesian skyline plot in BEAST, demographic events (population fluctuations) were further assessed over the history of Anatolian ground squirrels. Combined ecological niche modelling and molecular phylogeography revealed that S. xanthoprymnus, itself also a temperate (mid‐latitude) species, has responded to global climate changes through the Late Quaternary glacial–interglacial cycles in a fashion converse to that of most temperate (mid‐latitude) species: its range expanded rather than contracted during the glacial periods and contracted rather than expanded during the interglacial periods. In other words, Anatolian ground squirrels have been in refugia during the interglacial periods, suggesting that the classical paradigm of glacial range contraction and interglacial range expansion for temperate species may not be as general as previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 19–32.     

Phylogeny and comparative phylogeography of Sclerurus (Aves: Furnariidae) reveal constant and cryptic diversification in an old radiation of rain forest understorey specialists

Aim To evaluate the role of historical processes in the evolution of Sclerurus leaftossers by integrating phylogenetic and phylogeographical approaches. Location Humid forests of the Neotropical region. Methods We reconstructed the evolutionary history of Sclerurus based on DNA sequences representing all species and 20 of the 26 recognized subspecies using one autosomal nuclear locus and three protein‐coding mitochondrial gene sequences. Phylogenetic relationships were inferred using Bayesian and maximum‐likelihood methods. We used Bayesian coalescent‐based approaches to evaluate demographic changes through time, and to estimate the timing of diversification events. Based on these results, we examined the temporal accumulation of divergence events using lineage‐through‐time plots. Results The monophyly of all Sclerurus species was strongly supported except for Sclerurus mexicanus, which was paraphyletic in relation to Sclerurus rufigularis, and for the sister pair Sclerurus scansor–Sclerurus albigularis, which were not reciprocally monophyletic in the nuclear tree. We found remarkably deep phylogeographical structure within all Sclerurus species, and overall this structure was congruent with currently recognized subspecies and Neotropical areas of endemism. Diversification within Sclerurus has occurred at a relatively constant rate since the Middle Miocene. Main conclusions Our results strongly support the relevance of physiographical (e.g. Nicaragua Depression, Isthmus of Panama, Andean Cordillera, great rivers of Amazonia) and ecological barriers (open vegetation corridor) and ecological gradients (elevational zonation) to the diversification of Neotropical forest‐dwelling organisms. Despite the high congruence among the spatial patterns identified, the variance in divergence times suggests multiple speciation events occurring independently across the same barrier, and a role for dispersal. The phylogenetic patterns and cryptic diversity uncovered in this study demonstrate that the current taxonomy of Sclerurus underestimates the number of species.     

The genetic structure of crossbills suggests rapid diversification with little niche conservatism

Conservatism of ecological niches can cause geographical ranges or the formation of new species to be constrained, and might be expected in situations where strong trade‐offs result in ecological specialization. Here we address the flexibility of resource use in European crossbills by comparing the ecological and genetic similarities between four Mediterranean and three northern European crossbill populations, all specialized in feeding on a different resource. We used sequence data of one mitochondrial and two nuclear genes from between 211 and 256 individuals. The northern crossbills were genetically too similar to infer which population was more related to the southern ones. Crossbills from the island of Mallorca showed genetic signatures of a stable and isolated population, supporting their past treatment as a locally (co)evolving taxon, and seem to have evolved from an ecologically distinct ancestor. Previous studies in other populations also suggest that genetic similarity does not predict morphological and resource similarity. We estimate that the divergence of all western European crossbills has occurred within the last 11 000 years. Overall, it appears that crossbills can diversify rapidly and with little niche conservatism, but that such potentially reproductively isolated specialists are evolutionarily short‐lived. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 908–922.     

Fine‐scale group structure and demography of African savanna elephants recolonizing lands outside protected areas

Aim Dispersal is a critical component of animal ecology that is poorly understood for most species. In particular, savanna elephants (Loxodonta africana) have been studied for decades in national parks across Africa, but little is known about their dispersal into new or unused habitats or their population dynamics in human‐dominated landscapes. We capitalized on a natural dispersal event of savanna elephants recolonizing communal land in southern Kenya to document their demographic characteristics and genetic relationships. Location Rift Valley province of Kenya. Methods We collected faecal samples and used genetic methods to identify individuals, estimate the sex ratio and evaluate the patterns of relatedness within the female groups and male aggregations. We also measured dung bolus circumference to assign age classes to individuals and estimate the age structure. Results We identified 112 individuals with a sex ratio not different from one (1.32:1.00). The age structure was skewed towards younger elephants (71%), suggesting the potential for rapid growth from reproduction. We detected significantly higher kinship levels within female groups (R = 0.124 ± 0.023), suggesting that family groups colonized the site, but found little support for higher‐order genetic relationships among female groups. Males detected together were unrelated (R = 0.003 ± 0.030). Main conclusions Our results suggest that highly social mammals, such as savanna elephants, disperse into unoccupied habitat as family groups and that a young demographic structure and a large number of males might be expected in establishing populations. These findings highlight the potential value of indirect, non‐invasive methods for assessing elephant herd and demographic characteristics when direct observations are difficult.     

Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): species delimitation,morphological diagnosis,and refined mapping of geographical distribution

Although critical for enabling in‐depth evolutionary, ecological, or conservation‐orientated studies, taxonomic knowledge is still scarce for many groups of organisms, including mammals of the order Carnivora. For some of these taxa, even basic aspects such as species limits and geographical distribution are still uncertain. This is the case for the Neotropical mustelid genus Galictis, considered one of the least studied carnivoran genera in the Americas. To address this issue, we performed a comprehensive assessment of morphological and molecular characters to test the number of species within Galictis, and to characterize their distinctiveness and evolutionary history. In addition, we reviewed and consolidated the available information on the taxonomy of this genus, so as to provide a historical framework upon which we could interpret our data. Our analyses demonstrated that two Galictis species can be clearly delimited and diagnosed using metric and nonmetric morphological characters as well as DNA sequences from mitochondrial and nuclear gene segments. On the basis of this clarified species‐level delimitation, we reassessed the geographical range of each Galictis taxon, identifying possible areas of sympatry between them. These results provide a solid taxonomic framework for Galictis, enabling the development of additional studies focusing on this poorly known taxon. © 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167 , 449–472.     

Extending ecological niche models to the past 120 000 years corroborates the lack of strong phylogeographic structure in the Crested Drongo (Dicrurus forficatus forficatus) on Madagascar

We conduct a phylogeographic study of the Crested Drongo (Dicrurus forficatus forficatus), a broadly distributed bird species on Madagascar. We first determined the demographic and spatial pattern inferred from mitochondrial and nuclear data, and then compared these results with predictions from a present to 0.120‐Myr‐old reconstruction of the spatial dynamics of the range of D. f. forficatus on Madagascar, enabling putative areas of stability (lineage persistence) to be detected. Weak genetic structure along an east–west pattern and comparatively low genetic diversity were recovered, with strong evidence of population expansion found at all ten loci sampled. The palaeoclimatic distribution models over the past 0.120 Myr suggest the presence of extensive areas of suitable climate in the east and west for the species since its colonization of Madagascar, a result in strong concordance with the spatial and genetic signal derived from our multilocus data set. © 2013 The Linnean Society of London