Evolution of ontogenetic allometry shaping giant species: a case study from the damselfish genus Dascyllus (Pomacentridae)

The evolution of body size, the paired phenomena of giantism and dwarfism, has long been studied by biologists and paleontologists. However, detailed investigations devoted to the study of the evolution of ontogenetic patterns shaping giant species are scarce. The damselfishes of the genus Dascyllus appear as an excellent model for such a study. Their well understood phylogeny reveals that large‐bodied species have evolved in two different clades. Geometric morphometric methods were used to compare the ontogenetic trajectories of the neurocranium and the mandible in both small‐bodied (Dascyllus aruanus and Dascyllus carneus; maximum size: 50–65 mm standard length) and giant (Dascyllus trimaculatus and Dascyllus flavicaudus; maximum size: 90–110 mm standard length) Dascyllus species. At their respective maximum body size, the neurocranium of the giant species is significantly shorter and have a higher supraoccipital crest relative to the small‐bodied species, whereas mandible shape variation is more limited and is not related to the ‘giant’ trait. The hypothesis of ontogenetic scaling whereby the giant species evolved by extending the allometric trajectory of the small‐bodied ones (i.e. hypermorphosis) is rejected. Instead, the allometric trajectories vary among species by lateral transpositions. The rate of shape changes and the type of lateral transposition also differ according to the skeletal unit among Dascyllus species. Differences seen between the two giant species in the present study demonstrate that giant species may appear by varied alterations of the ancestor allometric pattern. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 99–117.     

Latitudinal clines in body size,but not in thermal tolerance or heat‐shock cognate 70 (HSC70), in the highly‐dispersing intertidal gastropod Littorina keenae (Gastropoda: Littorinidae)

Natural populations of widely‐distributed animals often exhibit clinal variation in phenotypic traits or in allele frequencies of a particular gene over their geographical range. A planktotrophic intertidal snail, Littorina keenae is broadly distributed along the north‐eastern Pacific coast through a large latitudinal range (24°50′N–43°18′N). We tested for latitudinal clines in two complex phenotypic traits – thermal tolerance and body size – and one single locus trait – heat shock cognate 70 (HSC70) – in L. keenae along almost its entire geographical range. We found only weak evidence for a latitudinal cline in the thermal tolerance and no evidence for a cline in allele frequencies at HSC70. However, as predicted by Bergmann's rule, we detected a strong latitudinal cline that accounted for 60% of the variance in body size (R² = 0.598; P < 0.001). In contrast, body size did not significantly affect thermal tolerance. HSC70 showed no genetic differentiation among the populations, supporting our previous mitochondrial gene‐based estimate of high gene flow during this snail's free‐swimming larval stage. Given that L. keenae experiences panmixia along its species range, the observed size cline may be partially or entirely caused by a phenotypically plastic response to local thermal environments rather than by genetic divergence in body size among populations in response to locally optimizing natural selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 494–505.     

Molecular data reveal that the tetraploid Tragopogon kashmirianus (Asteraceae: Lactuceae) is distinct from the North American T. mirus

Tragopogon kashmirianus (Asteraceae: Lactuceae) (2n = 24) was described based on collections from Kashmir. The tetraploid is morphologically similar to allotetraploid T. mirus from North America that has formed in western North America from the introduced T. dubius (2n = 12) and T. porrifolius (salsify; 2n = 12). Singh and Kachroo (1976 ) suggested that T. kashmirianus might have formed from the same diploid parental combination as T. mirus. To determine this, we investigated internal and external transcribed spacers (ITS, ETS) and five plastid regions of T. kashmirianus and species reported from Kashmir, northern India and neighbouring countries (T. badachschanicus, T. longirostris, T. porrifolius, T. pratensis, T. orientalis, T. subalpinus, T. trachycarpus, T. gracilis and T. dubius). Molecular data indicate that the parents of T. kashmirianus are not the European T. porrifolius and T. dubius. The exact parentage of T. kashmirianus is still unclear, but if it is an allotetraploid, at least one parent is a species native to Kashmir/India. Alternatively, it may represent an autopolyploid, again with the diploid parent native to Kashmir/India. We also found that ‘T. dubius’ from Kashmir is phylogenetically and morphologically distinct from collections of T. dubius from Europe and probably represents a previously unrecognized species. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 391–398.     

The role of size and colour pattern in protection of developmental stages of the red firebug (Pyrrhocoris apterus) against avian predators

We investigated how predator/prey body‐size ratio and prey colour pattern affected efficacy of prey warning signals. We used great and blue tits (Parus major and Cyanistes caeruleus), comprising closely related and ecologically similar bird species differing in body size, as experimental predators. Two larval instars and adults of the unpalatable red firebug (Pyrrhocoris apterus), differing in body size and/or coloration, were used as prey. We showed that prey body size did not influence whether a predator attacked the prey or not during the first encounter. However, smaller prey were attacked, killed, and eaten more frequently in repetitive encounters. We assumed that body size influences the predator through the amount of repellent chemicals better than through the amount of optical warning signal. The larger predator attacked, killed and ate all forms of firebug more often than the smaller one. The difference between both predators was more pronounced in less protected forms of firebug (chemically as well as optically). Colour pattern also substantially affected the willingness of predators to attack the prey. Larval red–black coloration did not provide a full‐value warning signal, although a similarly conspicuous red‐black coloration of the adults reliably protected them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 890–898.     

Coexistence of the predatory cyclopoids Acanthocyclops robustus (Sars) and Mesocyclops leuckarti (Claus) in a small eutrophic lake

The coexistence of two common species of predatory cyclopoid copepods, Acanthocyclops robustus and Mesocyclops leuckarti, in a small, eutrophic lake was studied. Both species are of similar size, inhabit the same water layers and have similar seasonal occurrence. A. robustus shows faster embryonic and postembryonic development and produces bigger clutches. Ingestion rates of A. robustus are higher than in M. leuckarti, especially with large prey types and low temperatures. Females of A. robustus are heavily preyed upon by fish. This predation pressure leads to extremely skewed sex ratios, which are unfavourable to population growth. A. robustus is therefore, on balance, favoured in productive lakes. Dominance over M. leuckarti is, however, less clear in warmer years.     

Bergmanns's size cline in New Zealand marine spray zone spiders (Araneae: Anyphaenidae: Amaurobioides)

Members of the spider genus Amaurobioides are restricted to the spray zone of rocky marine coasts, where they construct and hunt from silk retreats. Collecting for this study shows these spiders to be distributed around the entire New Zealand coast. A Templeton, Crandall, and Sing (TCS) analysis of the ND1 mitochondrial gene places specimens from the North Island and the northern half of the South Island into a group distinct from Amaurobioides maritima O.P.‐Cambridge, 1883, which is restricted to the southern half of the South Island. Females of this northern group exhibit latitude‐ and temperature‐related clines in body length, body mass, and residual index of condition, with larger individuals with greater indices of condition being found at cooler, southern sites. This size cline also appeared in a broader geographical analysis that included Amaurobioides piscator Hogg, 1909 from the sub‐Antarctic Auckland and Campbell Islands. Thirteen ND1 haplotypes are represented in the northern group. Both independent contrast analyses and standard regressions of the mean body lengths and mean masses of these haplotypes, and the mean latitudes and temperatures of the sites where haplotypes were present, document a Bergmann's size cline, and provide evidence for an underlying genetic component. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 78–92.     

The fellowship of the hobbit: the fauna surrounding Homo floresiensis

The Late Pleistocene Flores fauna shows a pattern observed on many other islands. It is neither aberrant nor exclusive, but the result of non‐random selective forces acting upon an impoverished and disharmonic insular fauna. By comparing the Flores vertebrate fauna with other fossil insular biotas, it is apparent that the evolution of Homo floresiensis is part of a general pattern affecting all the inhabitants of Pleistocene Flores. Vertebrate evolution on Flores appears to have been characterized by phylogenetic continuity, low species richness and a disharmonic fauna. All three aspects stem from the isolated position of the island and have resulted in the distinct morphological characteristics of the Flores fauna. Evidence reviewed herein shows that features exhibited by H. floresiensis, such as small stature, a small brain, relatively long arms, robust lower limbs and long feet, are not unique, but are shared by other insular taxa. Therefore, the evolution of H. floresiensis can be explained by existing models of insular evolution and followed evolutionary pathways similar to those of the other terrestrial vertebrates inhabiting Pleistocene Flores.     

Phylogeography of the Poecilimon luschani species group (Orthoptera,Tettigoniidae): a radiation strictly correlated with climatic transitions in the Pleistocene

The Quaternary biogeography of Anatolia has received considerable interest recently. Here, the genealogical history of the Anatolio?Balkan lineage of the Poecilimon luschani species group was evaluated. Using concatenated data from 16S rDNA and cytochrome c oxidase subunit I (COI) sequences, the timings of inter‐ and intraspecies radiations were estimated. The demographic history of the populations was estimated using a data set established from COI sequences. Genetic diversity was very high in almost all of the populations studied. Fixation indices suggested extreme divergence of P. luschani. A molecular chronogram estimated a radiation history for the species/subspecies over a period ranging from 1.323 to 0.440 Myr. Demographic analyses applied to 11 populations suggested departure in population size for most of the local populations. The following conclusions were reached: (1) P. luschani originated from an Anatolio‐Aegean ancestral stock and extended its range to the Balkans through Dardanelles during the Early Pleistocene; (2) the Mid‐Pleistocene Transition, the lengthening of glacial period from 41 to 100 Kyr and the initiation of intense glaciation periods are the three main events corresponding to the main nodes of the chronogram; (3) altitudinal heterogeneity played a buffer role during the glacial cycles, allowing populations to cope with severe environmental changes; (4) the effects of Pleistocene climate cycles on populations differ according to altitudinal and latitudinal location in Anatolia, and (5) habitat preferences, such as altitudinal range, may easily shift because of changes in environmental conditions. © 2014 The Linnean Society of London     

Factors affecting the vulnerability of cane toads (Bufo marinus) to predation by ants

Experimental evidence on the determinants of prey vulnerability is scarce, especially for vertebrates in the field. Invasive species offer robust opportunities to explore prey vulnerability, because the intensity of predation on or by such animals has not been eroded by coevolution. Around waterbodies in tropical Australia, native meat ants (Iridomyrmex reburrus) consume many metamorph cane toads (Bufo marinus, an invasive anuran). We document the determinants of toad vulnerability, especially the roles of toad body size and ant density. Larger metamorphs were attacked sooner (because they attracted more ants), but escaped more often. Overall, smaller toads were more likely to be killed. Ant densities influenced toad responses, as well as attack rate and success. Data on the immediate outcomes of attacks underestimate mortality: more than 73% of apparent ‘escapees’ died within 24 h. Because mortality during this period was independent of toad size, predation was less size selective than suggested by immediate outcomes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 738–751.     

Genetically and geographically isolated lineages of a tropical bat (Chiroptera: Molossidae) show demographic stability over the late Pleistocene

The newly described molossid bat, Chaerephon atsinanana Goodman et al., 2010, endemic to eastern Madagascar, shows notably high levels of phylogeographic and genetic structure compared with allopatric Chaerephon leucogaster Grandidier, 1869 from western Madagascar. Such highly significant structuring of haplotypes among altitudinally and latitudinally stratified population groups is contrary to the expected panmixia in strong flying bats. The null model of concordance in historical demographic patterns across these two Chaerephon species was not supported. Mismatch and Bayesian skyline analyses indicated ancient stable C. atsinanana populations of constant size during the last two major Pleistocene glacial periods, making retreat into and expansion from glacial refugia an unlikely explanation for such high levels of structure, in accordance with expectations for tropical bats. Analyses were consistent with post‐refugial population expansion in the less diverse and structured C. leucogaster during the end of the last Pleistocene glacial period. We hypothesise that the pronounced genetic structuring in C. atsinanana may result from female philopatry. Furthermore, differing demographic histories of the two species may have been shaped by differing climate or habitat preferences, consistent with evidence from MaxEnt ecological niche modelling, which shows differences in variables influencing the current predicted distributions. Fossil Quaternary pollen deposits further indicate greater stability in past climatic patterns in eastern versus western Madagascar. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 18–40.     

Phenotypic evolution in high‐elevation populations of western fence lizards (Sceloporus occidentalis) in the Sierra Nevada Mountains

Adaptive divergence in response to variable habitats, climates, and altitude is often accentuated along elevation gradients. We investigate phenotypic evolution in body size and coloration in the western fence lizard (Sceloporus occidentalis Baird & Girard, 1852) across elevation gradients in Yosemite National Park, California, situated in the Sierra Nevada mountains of Western North America. High‐elevation populations occurring above 2100 m a.s.l. are recognized as a separate subspecies (Sceloporus occidentalis taylori Camp, 1916), with a distinctive phenotype characterized by a large body size and extensive blue ventral pigmentation. We sampled S. occidentalis from across elevation gradients in Yosemite National Park, California, and collected phenotypic data (body size and ventral coloration measurements; 410 specimens) and mitochondrial DNA sequence data (complete NADH1 gene; 969 bp, 181 specimens) to infer phylogenetic relationships, and examine the genetic and phenotypic diversity among populations. Populations of S. occidentalis in Yosemite National Park follow Bergmann's rule and exhibit larger body sizes in colder, high‐elevation environments. The high‐elevation subspecies S. o. taylori is not monophyletic, and the mitochondrial DNA genealogy supports a model of convergent phenotypic evolution among high‐elevation populations belonging to different river drainages. The hypothesis that separate populations of S. occidentalis expanded up river drainages after the recession of glaciers is supported by population demographic analyses, and suggest that Bergmann's clines can evolve rapidly along elevation gradients. The distinctive high‐elevation phenotype that is attributable to S. o. taylori has evolved independently several times, and includes adaptive phenotypic changes associated with increases in body size and ventral coloration. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 630–641.     

A test of Rensch's rule in varanid lizards

In a model group of giant reptiles, we explored the allometric relationships between male and female body size and compared the effects of sexual and fecundity selection, as well as some proximate causes, on macroevolutionary patterns of sexual size dimorphism (SSD). Monitor lizards are a morphologically homogeneous group that has been affected by extreme changes in body size during their evolutionary history, resulting in 14‐fold differences among the body sizes of recent species. Here, we analysed data concerning the maximum and/or mean male and female snout–vent lengths in 42 species of monitor lizard from literary sources and supplemented these data with measurements made in zoos. There was a wide scale of SSD from nearly monomorphic species belonging mostly to the subgenus Odatria and Prasinus group of the Euprepriosaurus to apparently male‐larger taxa. The variable best explaining SSD was the body size itself; the larger the species, the higher the SSD. This pattern agrees with the currently discussed Rensch's rule, claiming that the relationship between male and female body size is hyperallometric, i.e. the allometric exponent of this relationship exceeds unity and thus SSD increases with body size in the case of male‐larger taxa. All our estimates of the reduced major axis regression slopes of this relationship ranged from 1.132 to 1.155. These estimates are significantly higher than unity, and thus unequivocally corroborate the validity of Rensch's rule in this reptilian group. In spite of our expectation that the variation in SSD can be alternatively explained by variables reflecting the strength of sexual selection (presence of male combat), fecundity selection (e.g. clutch size and mass) and/or proximate ecological factors (habitat type), none of these variables had consistent effects on SSD, especially when the data were adjusted to phylogenetic dependence and/or body size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 293–306.     

The early evolution of the mega‐diverse genus Begonia (Begoniaceae) inferred from organelle DNA phylogenies

Begonia L. is one of the largest flowering plant genera, a ubiquitous component of many tropical forests and an economically important ornamental plant. In the present study, we address the early evolution of Begonia by generating molecular phylogenies from approximately 7000 bases of chloroplast DNA and approximately 6000 bases of mitochondrial DNA for each of 30 exemplar Begonia species. Broadscale biogeographic patterns found in the phylogenies, together with previously estimated divergence dates, indicated that extant Begonia lineages first diversified in Africa and then subsequently in America and Asia. The phylogenies also revealed that the closest African relatives of the American and Asian Begonia are seasonally‐adapted species. Moderate to strong incongruence between the phylogenies suggested that they differ genealogically. These differences could have been the result of either interspecific hybridization and/or incomplete lineage sorting. The results obtained in the present study provide a much needed genus‐wide framework for future evolutionary studies of this exceptionally diverse tropical genus. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 243–250.     

Geographic variation in offspring size of a widespread lizard (Takydromus septentrionalis): importance of maternal investment

Geographic variation in offspring size is widespread, but the proximate causes of this variation have not yet been explicitly determined. We compared egg size and egg contents among five populations of a lizard (Takydromus septentrionalis, Günther, 1864) along a latitudinal gradient, and incubated eggs at two temperatures to determine the influence of maternal investment and incubation temperature on offspring size. The mean values for female size and egg size were both greater in the two northern populations (Chuzhou and Anji) than in the three southern populations (Lishui, Dongtou, and Ningde). The larger eggs were entirely attributable to the body size of females in the Anji population, but their increased size also stemmed from further enlargement of egg size relative to female body size in Chuzhou, the northernmost population sampled in this study. Eggs of the Chuzhou population contained more yolk and less water than those of southern populations. Despite the lower lipid content in the yolk, eggs from the Chuzhou population had higher energy contents than those from the two southern populations, owing to the larger egg size and increased volume of yolk. Hatchling size was not affected by incubation temperature, but differed significantly among populations, with hatchlings being larger in the Chuzhou population than in the other populations. Our data provide an inference that oviparous reptiles from cold climates may produce larger offspring, not only by increasing egg size but also by investing more energy into their eggs. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 59–67.     

The birth of an endemic species flock: demographic history of the Bellamya group (Gastropoda,Viviparidae) in Lake Malawi

Changes in habitat stability may significantly shape evolutionary patterns and processes in ancient lakes. In the present study, we use a hierarchical combination of molecular phylogenetic and coalescent approaches to investigate the evolutionary history of the endemic species of the gastropod genus Bellamya in the African rift‐lake Malawi. By integrating our findings with reported palaeontological and palaeolimnological data, we demonstrate that all but one evolutionary lineage of the Pliocene Bellamya fauna in Lake Malawi became extinct. Coalescent analyses indicate that the modern radiation underwent both a sudden demographic and a spatial expansion after a genetic bottleneck. We argue that a reflooding of the lake after severe Pleistocene low stands offers a straightforward explanation for this pattern and may have triggered speciation processes in the modern endemic Bellamya radiation in Lake Malawi. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 130–143.     

Historical demography of freshwater mussels (Bivalvia: Unionidae): genetic evidence for population expansion and contraction during the late Pleistocene and Holocene

Genetic variation was examined in two endangered mussel species, Epioblasma brevidens and Epioblasma capsaeformis, and in a non‐listed species, Lampsilis fasciola, in the Clinch River, Tennessee, USA, by screening mitochondrial DNA (mtDNA) sequences and nuclear DNA microsatellites. Patterns of mtDNA polymorphism exhibited different trends in long‐term population sizes for each species during the late Pleistocene and Holocene (～20 000 ya to present); namely, E. brevidens has declined over time, E. capsaeformis has remained demographically stable, and L. fasciola has expanded. However, analyses using microsatellites did not exhibit similar trends, perhaps because homoplasy had eliminated long‐term population signatures for the loci examined. For both marker types, long‐term effective population size (N_e) was low in E. brevidens, intermediate in E. capsaeformis, and high in L. fasciola. Moderately diverged mtDNA lineages, perhaps indicative of secondary contact, were observed in E. brevidens and E. capsaeformis. Perhaps the most surprising result of this study was the high level of genetic variation observed at both mtDNA and microsatellite DNA markers for L. fasciola, variation seemingly contrary to the relatively small demes that currently reside in the Clinch River. However, the data are consistent with known demographic and life‐history traits of these three mussel species and their fish hosts, namely that they each use hosts with different dispersal capabilities, ranging from low, moderate, and high, respectively. The low divergence of mtDNA sequence variation reported in this and other recent mussel studies indicates that considerable extant population genetic variation probably originated during the late Pleistocene and Holocene. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 376–397.     

Costs of secondary parasitism in the facultative hyperparasitoid Pachycrepoideus dubius: does host size matter?

Although hyperparasitism frequently occur in parasitic insects, many aspects of this strategy remain unknown. We investigated possible fitness costs of hyperparasitism as influenced by host size. Our study was conducted with the facultative hyperparasitoid Pachycrepoideus dubius Ashmead (Hymenoptera: Pteromalidae), which parasitizes host species differing greatly in size. We compared some fitness traits (level of successful parasitism, development time, sex ratio and offspring size) of P. dubius developing on large secondary/primary (Delia radicum L. (Diptera: Anthomyiidae)/Trybliographa rapae Westwood (Hymenoptera: Figitidae)) or small secondary/primary host species (Drosophila melanogaster L./Asobara tabida Nees (Hymenoptera: Braconidae)). In no-choice and choice experiments, P. dubius was able to develop on different stages of T. rapae (L2 (endophagous), L4 (ectophagous), and pupae) but that it preferred to parasitize unparasitized D. radicum pupae over pupae parasitized by T. rapae. Furthermore, in P. dubius, hyperparasitism was associated with fitness costs (lower level of successful parasitism, smaller adult size) and these costs were greater on the smallest host complex. We hypothesize that the size of D. melanogaster pupae parasitized by A. tabida may be close to the suboptimal host size for P. dubius beneath which the costs of hyperparasitism make this strategy nonadaptive. Hyperparasitism in terms of trade-offs between host quality and abundance of competitors is discussed.     

Comparative phylogeography reveals pre‐decline population structure of New Zealand Cyclodina (Reptilia: Scincidae) species

We examined the comparative phylogeography of all species within the endemic New Zealand skink genus Cyclodina to gain insight into the influence of historical processes on the biogeography of the North Island fauna. Until 1–2 kya, six Cyclodina species occurred sympatrically across the North Island of New Zealand. However, most species have undergone dramatic distributional declines subsequent to the introduction of mammals and the arrival of humans. We compare the phylogeographic patterns evident in Cyclodina species in three biogeographic categories: widespread species (Cyclodina aenea, Cyclodina ornata), North Island disjunct relics (Cyclodina macgregori, Cyclodina whitakeri), and northeastern island relics (Cyclodina alani, Cyclodina oliveri, Cyclodina townsi). Mitochondrial DNA (ND2) sequence data was obtained from across the entire range of each Cyclodina species. We used Neighbour‐joining, maximum likelihood and Bayesian methods to examine the phylogeographic patterns present in each species. Phylogeographic patterns varied among species in different biogeographic categories. Substantial phylogeographic structure was evident in the two widespread species (C. aenea, C. ornata), with Pliocene and Pleistocene divergences between clades evident. Divergences among island groups in the three northeastern island relic species (C. alani, C. oliveri, C. townsi) occurred during the late Pliocene–Pleistocene. By contrast, relatively shallow structure, indicative of late Pleistocene divergences, was present in the two North Island disjunct species (C. macgregori, C. whitakeri). The results strongly suggest that the Poor Knights Islands population of C. ornata represents a new species. We suggest that the contrasting phylogeographic patterns exhibited by Cyclodina species in different biogeographic categories might be related to body size, ecology, and habitat preferences. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 388–408.     

Ecophenotypic variation and phylogenetic inheritance in first lower molar shape of extant Italian populations of Microtus (Terricola) savii (Rodentia)

Seventeen extant populations of Microtus (Terricola) savii have been investigated for correlations of first lower molar shape to climatic variables by means of geometric morphometrics, and controlling for phylogenetic inheritance. Comparative methods revealed that climatic variables and phylogeography provide a very similar contribution to variation in first lower molars morphology, whereas tooth size does not appear to be affected by climatic conditions. Climate‐related changes have been recognized in the anteroconid portion of the tooth. This indicates that molar tooth variation is strongly influenced by climatic conditions, although in a complex way. Calabrian populations, often ascribed to Microtus (Terricola) brachycercus, form a distinct cluster, in agreement with the most recent genetic analyses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 632–647.     

Body mass and locomotor habits of the smallest darter,Anhinga minuta (Aves,Anhingidae)

During the Neogene of South America, Anhingidae was represented by several species, mainly with greater sizes than the extant members. In the present contribution, body mass and locomotor habits of Anhinga minuta, the smallest known darter, were inferred. Body mass was estimated using two methods, one with measures of a tibiotarsus (the holotype) and the other, with measurements of a humerus; locomotor habits were inferred through muscular reconstructions and wing parameters (wing span, wing area and wing loading). Estimates of wing span and wing area were based on the length of humerus, assuming a condition of isometry with respect to Anhinga anhinga; wing loading was obtained through a relation formula between wing area and body mass. The results obtained indicate a body mass of about 729 g, a wing span of 0.958 m, a wing area of 0.117 m² and a corresponding wing loading of 61 N/m². These values and also the proximal insertion of the musculus pectoralis are consistent with those of a soaring bird but with more frequent flapping than extant anhingids. Furthermore, the inferred musculature for tibiotarsus indicates abilities for swimming, climbing and moving through the vegetation as in extant representatives.