Biology Letters

Published by Royal Society, The
Online ISSN: 1744-957X
Publications
Humpback whale breeding stocks and suggested sub-stocks in the South Atlantic and western Indian Oceans with sighting locations for AHWC no. 1363 (asterisks). 
Humpback whale AHWC no. 1363 photographed (a) on Abrolhos Bank, Brazil and (b) off Madagascar. 
Article
Fidelity of individual animals to breeding sites is a primary determinant of population structure. The degree and scale of philopatry in a population reflect the fitness effects of social facilitation, ecological adaptation and optimal inbreeding. Patterns of breeding-site movement and fidelity are functions of social structure and are frequently sex biased. We report on a female humpback whale (Megaptera novaeangliae) first identified by natural markings off Brazil that subsequently was photographed off Madagascar. The minimum travel distance between these locations is greater than 9800 km, approximately 4000 km longer than any previously reported movement between breeding grounds, more than twice the species' typical seasonal migratory distance and the longest documented movement by a mammal. It is unexpected to find this exceptional long-distance movement between breeding groups by a female, as models of philopatry suggest that male mammals move more frequently or over longer distances in search of mating opportunities. While such movement may be advantageous, especially in changeable or unpredictable circumstances, it is not possible to unambiguously ascribe causality to this rare observation. This finding illustrates the behavioural flexibility in movement patterns that may be demonstrated within a typically philopatric species.
 
Box plots of intraspecific divergence observed for populations of species that were collected from two or more localities. The median distances of separation are given for the general localities of Churchill, MB (C), Guelph, ON (G), Ottawa, ON (O), St Andrew's, NB (N) and Great Smoky Mountains National Park, TN/NC (T). The number of comparisons used to calculate genetic divergence is denoted above each box plot.
Histograms and neighbour-joining trees showing deep sequence divergences at COI among species in three genera of Lepidoptera from eastern North America: (a) Plusia, (b) Hyphantria and (c) Spodoptera. For the species displaying deep divergences, intraspecific divergences are shaded red and divergences among congeneric taxa are shaded blue. Individuals showing deep ‘intraspecific’ barcode divergence occur in sympatry for all three taxa.
Article
This study reports DNA barcodes for more than 1300 Lepidoptera species from the eastern half of North America, establishing that 99.3 per cent of these species possess diagnostic barcode sequences. Intraspecific divergences averaged just 0.43 per cent among this assemblage, but most values were lower. The mean was elevated by deep barcode divergences (greater than 2%) in 5.1 per cent of the species, often involving the sympatric occurrence of two barcode clusters. A few of these cases have been analysed in detail, revealing species overlooked by the current taxonomic system. This study also provided a large-scale test of the extent of regional divergence in barcode sequences, indicating that geographical differentiation in the Lepidoptera of eastern North America is small, even when comparisons involve populations as much as 2800 km apart. The present results affirm that a highly effective system for the identification of Lepidoptera in this region can be built with few records per species because of the limited intra-specific variation. As most terrestrial and marine taxa are likely to possess a similar pattern of population structure, an effective DNA-based identification system can be developed with modest effort.
 
Article
We present the first genomic-scale analysis addressing the phylogenetic position of turtles, using over 1000 loci from representatives of all major reptile lineages including tuatara. Previously, studies of morphological traits positioned turtles either at the base of the reptile tree or with lizards, snakes and tuatara (lepidosaurs), whereas molecular analyses typically allied turtles with crocodiles and birds (archosaurs). A recent analysis of shared microRNA families found that turtles are more closely related to lepidosaurs. To test this hypothesis with data from many single-copy nuclear loci dispersed throughout the genome, we used sequence capture, high-throughput sequencing and published genomes to obtain sequences from 1145 ultraconserved elements (UCEs) and their variable flanking DNA. The resulting phylogeny provides overwhelming support for the hypothesis that turtles evolved from a common ancestor of birds and crocodilians, rejecting the hypothesized relationship between turtles and lepidosaurs.
 
Bayesian tree for Charadriiformes genera. Letters A to P indicate nodes for which fossil or molecular time constraints were used to estimate divergence times (see table 2 in electronic supplementary material). Numbers at nodes are posterior probabilities (PP), which are not indicated if PPZ1.0. Nodes with PP!0.5 are collapsed.
Bayesian chronogram for Charadriiformes genera. The Cretaceous/Paleocene boundary is marked by a grey bar. Nodes are labelled A, C, L and S for Avian outgroups, Charadrii, Lari and Scolopaci divergences, respectively.  
Article
Comparative study of character evolution in the shorebirds is presently limited because the phylogenetic placement of some enigmatic genera remains unclear. We therefore used Bayesian methods to obtain a well-supported phylogeny of 90 recognized genera using 5 kb of mitochondrial and nuclear sequences. The tree comprised three major clades: Lari (gulls, auks and allies plus buttonquails) as sister to Scolopaci (sandpipers, jacanas and allies), and in turn sister to Charadrii (plovers, oystercatchers and allies), as in previous molecular studies. Plovers and noddies were not recovered as monophyletic assemblages, and the Egyptian plover Pluvianus is apparently not a plover. Molecular dating using multiple fossil constraints suggests that the three suborders originated in the late Cretaceous between 79 and 102 Mya, and at least 14 lineages of modern shorebirds survived the mass extinction at the K/T boundary. Previous difficulties in determining the phylogenetic relationships of enigmatic taxa reflect the fact that they are well-differentiated relicts of old, genus-poor lineages. We refrain from suggesting systematic revisions for shorebirds at this time because gene trees may fail to recover the species tree when long branches are connected to deep, shorter branches, as is the case for some of the enigmatic taxa.
 
Article
The conference 'Celebrating Darwin: From the Origin of Species to Deep Metazoan Phylogeny' was held at the Humboldt University in Berlin, from 3 to 6 March 2009. Specialists from the fields of bioinformatics, molecular biology, developmental biology, comparative morphology and paleontology joined forces to present and discuss novel approaches in reconstructing the still unresolved early branching patterns of the metazoan tree of life.
 
(a) Map of the study area. Stippled areas depict floodplain and unfilled areas savannah woodland. Filled circles denote lizards displaying Fogg Dam ND2-ND4 haplotypes and open circles denote lizards with Scotts Creek ND2-ND4 haplotypes. Lizard M5 is depicted as a mix of the two haplotypes. (b) Unrooted tree based on Bayesian analysis of concatenated mitochondrial ND4 and ND2 genes. 
Article
Mitochondrial DNA (mtDNA) is the traditional workhorse for reconstructing evolutionary events. The frequent use of mtDNA in such analyses derives from the apparent simplicity of its inheritance: maternal and lacking bi-parental recombination. However, in hybrid zones, the reproductive barriers are often not completely developed, resulting in the breakdown of male mitochondrial elimination mechanisms, leading to leakage of paternal mitochondria and transient heteroplasmy, resulting in an increased possibility of recombination. Despite the widespread occurrence of heteroplasmy and the presence of the molecular machinery necessary for recombination, we know of no documented example of recombination of mtDNA in any terrestrial wild vertebrate population. By sequencing the entire mitochondrial genome (16761bp), we present evidence for mitochondrial recombination in the hybrid zone of two mitochondrial haplotypes in the Australian frillneck lizard (Chlamydosaurus kingii).
 
Article
Parenting is one of the main influences on children's early development, and yet its underlying genetic mechanisms have only recently begun to be explored, with many studies neglecting to control for possible child effects. This study focuses on maternal behaviour and on an allele at the RS3 promoter region of the arginine vasopressin receptor 1A (AVPR1A) gene, previously associated with autism and with higher amygdala activation in a face-matching task. Mothers were observed during a free-play session with each of their 3.5-year-old twins. Multilevel regression analyses revealed that mothers who are carriers of the AVPR1A RS3 allele tend to show less structuring and support throughout the interaction independent of the child's sex and RS3 genotype. This finding advances our understanding of the genetic influences on human maternal behaviour.
 
Enamel d 13 C and d 18 O values for the fossil ( a ) bovid and ( b ) suid specimens sampled. These are differentiated by stratigraphic level within the Shungura Formation. Specimens from units C-4 and younger (i.e. , 2.8 Ma) are significantly enriched in d 13 C relative to those from units B-10 and older. 
Article
Late Pliocene climate changes have long been implicated in environmental changes and mammalian evolution in Africa, but high-resolution examinations of the fossil and climatic records have been hampered by poor sampling. By using fossils from the well-dated Shungura Formation (lower Omo Valley, northern Turkana Basin, southern Ethiopia), we investigate palaeodietary changes in one bovid and in one suid lineage from 3 to 2 Ma using stable isotope analysis of tooth enamel. Results show unexpectedly large increases in C(4) dietary intake around 2.8 Ma in both the bovid and suid, and possibly in a previously reported hippopotamid species. Enamel δ(13)C values after 2.8 Ma in the bovid (Tragelaphus nakuae) are higher than recorded for any living tragelaphin, and are not expected given its conservative dental morphology. A shift towards increased C(4) feeding at 2.8 Ma in the suid (Kolpochoerus limnetes) appears similarly decoupled from a well-documented record of dental evolution indicating gradual and progressive dietary change. The fact that two, perhaps three, disparate Pliocene herbivore lineages exhibit similar, and contemporaneous changes in dietary behaviour suggests a common environmental driver. Local and regional pollen, palaeosol and faunal records indicate increased aridity but no corresponding large and rapid expansion of grasslands in the Turkana Basin at 2.8 Ma. Our results provide new evidence supporting ecological change in the eastern African record around 2.8 Ma, but raise questions about the resolution at which different ecological proxies may be comparable, the correlation of vegetation and faunal change, and the interpretation of low δ(13)C values in the African Pliocene.
 
Article
Analysis of DNA sequences now plays a key role in evolutionary biology research. If Darwin were to come back today, I think he would be absolutely delighted with molecular evolutionary genetics, for three reasons. First, it solved one of the greatest problems for his theory of evolution by natural selection. Second, it gives us a tool that can be used to investigate many of the questions he found the most fascinating. And third, DNA data confirm Darwin's grand view of evolution.
 
Article
Although the hymenopteran sex-determining mechanism generally results in haploid males and diploid females, diploid males can be produced via homozygosity at the sex-determining locus. Diploid males have low fitness because they are effectively sterile or produce presumably sterile triploid offspring. Previously, triploid females were observed in three species of North American Polistes paper wasps, and this was interpreted as indirect evidence of diploid males. Here we report what is, to our knowledge, the first direct evidence: four of five early male-producing Polistes dominulus nests from three populations contained diploid males. Because haploid males were also found, however, the adaptive value of early males cannot be ignored. Using genetic and morphological data from triploid females, we also present evidence that both diploid males and triploid females remain undetected throughout the colony cycle. Consequently, diploid male production may result in a delayed fitness cost for two generations. This phenomenon is particularly relevant for introduced populations with few alleles at the sex-determining locus, but cannot be ignored in native populations without supporting genetic data. Future research using paper wasp populations to test theories of social evolution should explicitly consider the potential impacts of diploid males.
 
Article
The concept of 'evolvability' is increasingly coming to dominate considerations of evolutionary change. There are, however, a number of different interpretations that have been put on the idea of evolvability, differing in the time scales over which the concept is applied. For some, evolvability characterizes the potential for future adaptive mutation and evolution. Others use evolvability to capture the nature of genetic variation as it exists in populations, particularly in terms of the genetic covariances between traits. In the latter use of the term, the applicability of the idea of evolvability as a measure of population's capacity to respond to natural selection rests on one, but not the only, view of the way in which we should envisage the process of natural selection. Perhaps the most potentially confusing aspects of the concept of evolvability are seen in the relationship between evolvability and robustness.
 
Article
Times for indoor 200 m sprint races are notably worse than those for outdoor races. In addition, there is a considerable bias against competitors drawn in inside lanes (with smaller bend radii). Centripetal acceleration requirements increase average forces during sprinting around bends. These increased forces can be modulated by changes in duty factor (the proportion of stride the limb is in contact with the ground). If duty factor is increased to keep limb forces constant, and protraction time and distance travelled during stance are unchanging, bend-running speeds are reduced. Here, we use results from the 2004 Olympics and World Indoor Championships to show quantitatively that the decreased performances in indoor competition, and the bias by lane number, are consistent with this 'constant limb force' hypothesis. Even elite athletes appear constrained by limb forces.
 
Article
To Darwin, parasites were fascinating examples of adaptation but their significance as selective factors for a wide range of phenomena has only been studied in depth over the last few decades. This work has had its roots in behavioural/evolutionary ecology on the one hand, and in population biology/ecology on the other, thus shaping a new comprehensive field of 'evolutionary parasitology'. Taking parasites into account has been a success story and has shed new light on several old questions such as sexual selection, the evolution of sex and recombination, changes in behaviour, adaptive life histories, and so forth. In the process, the topic of ecological immunology has emerged, which analyses immune defences in a framework of costs and benefits. Throughout, a recurrent theme is how to appropriately integrate the underlying mechanisms as evolved boundary conditions into a framework of studying the adaptive value of traits. On the conceptual side, major questions remain and await further study.
 
Article
Darwin devoted much of his working life to the study of plant reproductive systems. He recognized that many of the intricacies of floral morphology had been shaped by natural selection in favour of outcrossing, and he clearly established the deleterious effects of self-fertilization on progeny. Although Darwin hypothesized the adaptive significance of self-fertilization under conditions of low mate availability, he held that a strategy of pure selfing would be strongly disadvantageous in the long term. Here, I briefly review these contributions to our understanding of plant reproduction. I then suggest that investigating two very different sexual systems, one in plants and the other in animals, would throw further light on the long-term implications of a commitment to reproduction exclusively by selfing.
 
Characteristics of eusocial taxa. taxon sex determination type of development nesting habits worker characteristics food source 
Article
Darwin identified eusocial evolution, especially of complex insect societies, as a particular challenge to his theory of natural selection. A century later, Hamilton provided a framework for selection on inclusive fitness. Hamilton's rule is robust and fertile, having generated multiple subdisciplines over the past 45 years. His suggestion that eusociality can be explained via kin selection, however, remains contentious. I review the continuing debate on the role of kin selection in eusocial evolution and suggest some lines of research that should resolve that debate.
 
Numbers of case and control reports available from each of the RBCT triplets for the various analyses. 
Logistic regression model of the risk of a TB breakdown showing the prevalence of exposure to the various factors among cases and population attributable fractions. 
Article
A case-control study of the factors associated with the risk of a bovine tuberculosis (TB) breakdown in cattle herds was undertaken within the randomized badger culling trial (RBCT). TB breakdowns occurring prior to the 2001 foot-and-mouth disease epidemic in three RBCT triplets were eligible to be cases; controls were selected from the same RBCT area. Data from 151 case farms and 117 control farms were analysed using logistic regression. The strongest factors associated with an increased TB risk were movement of cattle onto the farm from markets or farm sales, operating a farm over multiple premises and the use of either covered yard or 'other' housing types. Spreading artificial fertilizers or farmyard manure on grazing land were both associated with decreased risk. These first case-control results from the RBCT will be followed by similar analyses as more data become available.
 
Article
[Mitchell & Lust (2008)][1] present evidence that the carotid rete of artiodactyls allows for a dissociation between brain and carotid artery temperatures, muting whole body thermoregulatory responses, and thus allowing for energy and water conservation. They argue that this morphological feature
 
Article
Biomechanics Comment There is always a trade-off between speed and force in a lever system: comment on McHenry (2010) In a recent Biology Letters article, McHenry [1] makes a distinction between levers that operate under 'quasi-static' and 'dynamic' conditions, concluding that 'no trade-off between force and velocity exists in a lever with spring – mass dynamics'. As evidence, McHenry uses a computer model to simulate a kicking locust leg powered by a spring. While we concur with aspects of McHenry's analysis—and we agree that complex relationships between forces and speeds may emerge from some biological levers—we offer a different interpretation of McHenry's data.
 
Article
Leal & Powell [[1][1]] (L&P) report experiments on the tropical lizard Anolis evermanni which, in their view, ‘show that A. evermanni exhibits behavioural flexibility across multiple cognitive tasks, including solving a novel motor task using multiple strategies and reversal learning, plus rapid
 
Article
Galhardo et al . [[1][1]] present the results of an experiment designed to test how the social environment modulates the expression of two personality traits, exploration–avoidance and neophobia. The authors highlight that novel object (NO) tests are frequently used to measure the personality
 
Article
Ecosystems may respond to changes in environmental conditions by abruptly switching between alternative stable states [[1][1]], yet the mechanisms underlying these changes are poorly understood. In the North Sea, the distribution and abundance of species across several trophic levels have changed
 
( a ) The change in the dinosaur body-size distribution in the Dinosaur Park Formation through successive collecting years. ( b ) Reduction in negative skew of the body-size distribution as sampling increases through successive collection years. 
Article
Codron et al. [[1][1]] invoke an ecological model of size-specific competition in dinosaurs to explain an apparent bimodal distribution within Dinosauria, and find ‘intermediate-sized taxa’ (1–1000 kg) are prone to extinction. Although the authors take an interesting approach, we argue that
 
Article
Are the associations between the facial width-to-height ratio (fWHR) and Japanese male baseball player performance documented by Tsujimura & Banissy [[1][1]] robust to controlling for body mass index (BMI)? Two factors motivate this question. First, research suggests that BMI is positively
 
(a) An adult green hermit (P. guy; inset) and the same species with a transmitter attached for monitoring movements. (b) An example of two hummingbird movement paths through agricultural landscapes. Movement paths are shown in yellow with telemetry locations in blue. Direct-line distances between release ( blue triangles) and capture locations (green squares) are shown in black. Forested areas are in green, agriculture in brown. Photo credits: J. Miller (a), M. Betts (b). 
Article
Reduced pollination success, as a function of habitat loss and fragmentation, appears to be a global phenomenon. Disruption of pollinator movement is one hypothesis put forward to explain this pattern in pollen limitation. However, the small size of pollinators makes them very difficult to track; thus, knowledge of their movements is largely speculative. Using tiny radio transmitters (0.25 g), we translocated a generalist tropical 'trap-lining' hummingbird, the green hermit (Phaethornis guy), across agricultural and forested landscapes to test the hypothesis that movement is influenced by patterns of deforestation. Although, we found no difference in homing times between landscape types, return paths were on average 459+/-144 m (+/-s.e.) more direct in forested than agricultural landscapes. In addition, movement paths in agricultural landscapes contained 36+/-4 per cent more forest than the most direct route. Our findings suggest that this species can circumvent agricultural matrix to move among forest patches. Nevertheless, it is clear that movement of even a highly mobile species is strongly influenced by landscape disturbance. Maintaining landscape connectivity with forest corridors may be important for enhancing movement, and thus in facilitating pollen transfer.
 
Article
Distributions of mutation fitness effects from evolution experiments are available in an increasing number of species, opening the way for a vast array of applications in evolutionary biology. However, comparison of estimated distributions among studies is hampered by inconsistencies in the definitions of fitness effects and selection coefficients. In particular, the use of ratios of Malthusian growth rates as 'relative fitnesses' leads to wrong inference of the strength of selection. Scaling Malthusian fitness by the generation time may help overcome this shortcoming, and allow accurate comparison of selection coefficients across species. For species reproducing by binary fission (neglecting cellular death), ln2 can be used as a correction factor, but in general, the growth rate and generation time of the wild-type should be provided in studies reporting distribution of mutation fitness effects. I also discuss how density and frequency dependence of population growth affect selection and its measurement in evolution experiments.
 
Article
The temporal stability of plant reproductive features on islands has rarely been tested. Using flowers, fruits/cones and seeds from a well-dated (23 Ma) Miocene Lagerstätte in New Zealand, we show that across 23 families and 30 genera of forest angiosperms and conifers, reproductive features have remained constant for more than 20 Myr. Insect-, wind- and bird-pollinated flowers and wind- and bird-dispersed diaspores all indicate remarkable reproductive niche conservatism, despite widespread environmental and biotic change. In the past 10 Myr, declining temperatures and the absence of low-latitude refugia caused regional extinction of thermophiles, while orogenic processes steepened temperature, precipitation and nutrient gradients, limiting forest niches. Despite these changes, the palaeontological record provides empirical support for evidence from phylogeographical studies of strong niche conservatism within lineages and biomes.
 
(a) Simultaneous reciprocal copulation of Bradybaena similaris (BS) and (b) penial removal by Bradybaena pellucida (BP) from BS in the middle of copulation. Arrows indicate the everted penes. In (b), the BP penis is no longer connected to the BS snail. 
Asymmetry of reproductive isolation during simultaneous reciprocal mating between B. pellucida (BP) and B. similaris (BS). (a) Proportions of ovipositing individuals. (b) Proportions of individuals that received a spermatophore. (c) Copulation duration. Open and filled bars indicate the means (Gs.e.) for BP and BS, respectively. Values in parentheses indicate the numbers of individuals. 
Article
The generality of asymmetric reproductive isolation between reciprocal crosses suggests that the evolution of isolation mechanisms often proceeds in reciprocal asymmetry. In hermaphroditic snails that copulate simultaneously and reciprocally, asymmetry in premating isolation may not be readily detectable because the failure of the symmetric performance of courtship would prevent copulation from occurring. On the other hand, through their prolonged copulation, snails discriminate among mates when exchanging spermatophores for their benefit and thus may exhibit asymmetric reproductive isolation during interspecific mating. However, no clear case of reciprocal asymmetry has been found in reproductive isolation between snail species. Here we show a discrete difference in hybridization success between simultaneous reciprocal copulations between two species of pulmonate snails. Premating isolation of Bradybaena pellucida (BP) and Bradybaena similaris (BS) is incomplete in captivity. In interspecific copulation, BP removes its penis without transferring a spermatophore, while BS sires hybrids by inseminating BP. Thus, 'male' BP or 'female' BS rejects the other individual, while female BP and male BS accept each other, so that the two sexes of either BP or BS oppose each other in mate discrimination. Our results are a clear example of asymmetry in reproductive isolation during simultaneous reciprocal mating between hermaphroditic animals.
 
Mean growth ( G s.e.) of A. polyacanthus juveniles on high (solid lines) and low (dashed lines) quantity diets, and in relation to the feeding regime of their parents on high (black circles) or low (white circles) food diet. 
Mean survival ( G s.e.) of A. polyacanthus offspring produced by parents consuming high (filled circles) and low (open circles) food diets in ( a ) juvenile in high-food environments and ( b ) juvenile in low-food environments. 
Article
Both the parental legacy and current environmental conditions can affect offspring life histories; however, their relative importance and the potential relationship between these two influences have rarely been investigated. We tested for the interacting effects of parental and juvenile environments on the early life history of the marine fish Acanthochromis polyacanthus. Juveniles from parents in good condition were longer and heavier at hatching than juveniles from parents in poor condition. Parental effects on juvenile size were evident up to 29 days post-hatching, but disappeared by 50 days. Offspring from good condition parents had higher early survival than offspring from poor-condition parents when reared in a low-food environment. By contrast, parental condition did not affect juvenile survival in the high-food environment. These results suggest that parental effects on offspring performance are most important when poor environmental conditions are encountered by juveniles. Furthermore, parental effects observed at hatching may often be moderated by compensatory mechanisms when environmental conditions are good.
 
(a) The apparatus used for artificial cyclic bending of feathers; (b) the testing set-up.  
Bending stiffness (P/d [N mm K1 ], i.e. external load divided by displacement) as a function of fatigue duration (h). (a) Willow warbler (Phylloscopus trochilus); (b) chiffchaff (Phylloscopus collybita). Different symbols represent individual feathers and lines are regression lines.  
Article
Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.
 
Genetic map of the thelytoky region. Marker names are displayed on the right, genetic distances (cM) between markers are given on the left. The graph shows the LOD scores for two-point comparisons of the respective marker and the thelytoky gene.  
Article
Differentiation into castes and reproductive division of labour are a characteristics of eusocial insects. Caste determination occurs at an early stage of larval development in social bees and is achieved via differential nutrition irrespective of the genotype. Workers are usually subordinate to the queen and altruistically refrain from reproduction. Workers of the Cape honeybee (Apis mellifera capensis) do not necessarily refrain from reproduction. They have the unique ability to produce female offspring parthenogenetically (thelytoky) and can develop into 'pseudoqueens'. Although these are morphologically workers, they develop a queen-like phenotype with respect to physiology and behaviour. Thelytoky is determined by a single gene (th) and we show that this gene also influences other traits related to the queen phenotype, including egg production and queen pheromone synthesis. Using 566 microsatellite markers, we mapped this gene to chromosome 13 and identified a candidate locus thelytoky, similar to grainy head (a transcription factor), which has been shown to be highly expressed in queens of eusocial insects. We therefore suggest that this gene is not only important for determining the pseudoqueen phenotype in A. m. capensis workers, but is also of general importance in regulating the gene cascades controlling reproduction and sterility in female social bees.
 
Scatterplot of the length of the fingers on right hands in 297 biology students. The common line estimated by the ordinary least-square linear regression is depicted as the solid line (length of the 2nd finger ¼ 0.823 (length of the 4th finger) þ 11.223; p , 0.00001; r 2 ¼ 0.81). Its intercept is significantly larger than zero (b ¼ 11.223 + 1.654; p , 0.00001), which means the 2D : 4D ratio necessarily decreases with increasing finger length. The sexes do not significantly differ in the scaling relationship between the length of the 2nd and 4th finger (see table 1 for statistics). Women, circles and dotted line; men, squares and broken line.  
Article
Ratios often lead to biased conclusions concerning the actual relationships between examined traits and comparisons of the relative size of traits among groups. Therefore, the use of ratios has been abandoned in most comparative studies. However, ratios such as body mass index and waist-to-hip ratio are widely used in evolutionary biology and medicine. One such, the ratio of the 2nd to the 4th finger (2D : 4D), has been the subject of much recent interest in both humans and animals. Most studies agree that 2D : 4D is sexually dimorphic. In men, the 2nd digit tends to be shorter than the 4th, while in women the 2nd digit tends to be of the same size or slightly longer than the 4th. Nevertheless, here we demonstrate that the sexes do not greatly differ in the scaling between the 2nd and 4th digit. Sexual differences in 2D : 4D are mainly caused by the shift along the common allometric line with non-zero intercept, which means 2D : 4D necessarily decreases with increasing finger length, and the fact that men have longer fingers than women. We conclude that previously published results on the 2D : 4D ratio are biased by its covariation with finger length. We strongly recommend regression-based approaches for comparisons of hand shape among different groups.
 
Multilocus heterozygosity correlates with T-lymphocyte proliferation in response to phytohaemagglutinin (PHA) injection. We calculated PHA response as the ratio of post-injection (48 h) patagial width to pre-injection width. 
Article
Evidence is accumulating that genetic variation within individual hosts can influence their susceptibility to pathogens. However, there have been few opportunities to experimentally test this relationship, particularly within outbred populations of non-domestic vertebrates. We performed a standardized pathogen challenge in house finches (Carpodacus mexicanus) to test whether multilocus heterozygosity across 12 microsatellite loci predicts resistance to a recently emerged strain of the bacterial pathogen, Mycoplasma gallisepticum (MG). We simultaneously tested whether the relationship between heterozygosity and pathogen susceptibility is mediated by differences in cell-mediated or humoral immunocompetence. We inoculated 40 house finches with MG under identical conditions and assayed both humoral and cell-mediated components of the immune response. Heterozygous house finches developed less severe disease when infected with MG, and they mounted stronger cell-mediated immune responses to phytohaemagglutinin. Differences in cell-mediated immunocompetence may, therefore, partly explain why more heterozygous house finches show greater resistance to MG. Overall, our results underscore the importance of multilocus heterozygosity for individual pathogen resistance and immunity.
 
Article
Fig wasps and fig trees are mutually dependent, with each of the 800 or so species of fig trees (Ficus, Moraceae) typically pollinated by a single species of fig wasp (Hymenoptera: Agaonidae). Molecular evidence suggests that the relationship existed over 65 Ma, during the Cretaceous. Here, we record the discovery of the oldest known fossil fig wasps, from England, dated at 34 Ma. They possess pollen pockets that contain fossil Ficus pollen. The length of their ovipositors indicates that their host trees had a dioecious breeding system. Confocal microscopy and scanning electron microscopy reveal that the fossil female fig wasps, and more recent species from Miocene Dominican amber, display the same suite of anatomical characters associated with fig entry and pollen-carrying as modern species. The pollen is also typical of modern Ficus. No innovations in the relationship are discernible for the last tens of millions of years.
 
Article
The extreme body size of blue whales requires a high energy intake and therefore demands efficient foraging strategies. As an obligate lunge feeder on aggregations of small zooplankton, blue whales engulf a large volume of prey-laden water in a single, rapid gulp. The efficiency of this feeding mechanism is strongly dependent on the amount of prey that can be captured during each lunge, yet food resources tend to be patchily distributed in both space and time. Here, we measured the three-dimensional kinematics and foraging behaviour of blue whales feeding on krill, using suction-cup attached multi-sensor tags. Our analyses revealed 360° rolling lunge-feeding manoeuvres that reorient the body and position the lower jaws so that a krill patch can be engulfed with the whale's body inverted. We also recorded these rolling behaviours when whales were in a searching mode in between lunges, suggesting that this behaviour also enables the whale to visually process the prey field and maximize foraging efficiency by surveying for the densest prey aggregations. These results reveal the complex manoeuvrability that is required for large rorqual whales to exploit prey patches and highlight the need to fully understand the three-dimensional interactions between predator and prey in the natural environment.
 
Bayesian unrooted phylogeny of eukaryotes, with a basal trichotomy representing uncertainties in the relationships between the three groups. The tree was obtained from the consensus between two independent Markov chains, run under the CAT model implemented in PHYLOBAYES. The species colour code corresponds to the type of plastid pigments, as follows: purple, chlorophyll a; green, chlorophyll aCb; and red, chlorophyll aCc. The asterisks represent primary, secondary or tertiary endosymbiosis. Underlined numbers at nodes represent PP of the analysis performed with the constant sites removed/analysis performed with all sites; other numbers represent the result of the ML bootstrap analysis (BS)-Node 1 below the line: ML analysis of the full-length alignment//ML analysis with category 7 removed/ML analysis with catergories 6C7 removed. Black dots correspond to 1.0 PP and 100% BS; black squares correspond to 1.0 PP and the specified values of BS. The scale bar represents the estimated number of amino acid substitutions per site. 
Article
Advances in molecular phylogeny of eukaryotes have suggested a tree composed of a small number of supergroups. Phylogenomics recently established the relationships between some of these large assemblages, yet the deepest nodes are still unresolved. Here, we investigate early evolution among the major eukaryotic supergroups using the broadest multigene dataset to date (65 species, 135 genes). Our analyses provide strong support for the clustering of plants, chromalveolates, rhizarians, haptophytes and cryptomonads, thus linking nearly all photosynthetic lineages and raising the question of a possible unique origin of plastids. At its deepest level, the tree of eukaryotes now receives strong support for two monophyletic megagroups comprising most of the eukaryotic diversity.
 
Article
Populations of sturgeon (Acipenseridae) have experienced global declines, and in some cases extirpation, during the past century. In the current era of climate change and over-harvesting of fishery resources, climate models, based on uncertain boundary conditions, are being used to predict future effects on the Earth's biota. A collection of approximately 400-year-old Atlantic sturgeon spines from a midden in colonial Jamestown, VA, USA, allowed us to compare the age structure and growth rate for a pre-industrial population during a 'mini-ice age' with samples collected from the modern population in the same reach of the James River. Compared with modern fish, the colonial population was characterized by larger and older individuals and exhibited significantly slower growth rates, which were comparable with modern populations at higher latitudes of North America. These results may relate to higher population densities and/or colder water temperatures during colonial times.
 
Article
Despite hopes that the processes of molecular evolution would be simple, clock-like and essentially universal, variation in the rate of molecular evolution is manifest at all levels of biological organization. Furthermore, it has become clear that rate variation has a systematic component: rate of molecular evolution can vary consistently with species body size, population dynamics, lifestyle and location. This suggests that the rate of molecular evolution should be considered part of life-history variation between species, which must be taken into account when interpreting DNA sequence differences between lineages. Uncovering the causes and correlates of rate variation may allow the development of new biologically motivated models of molecular evolution that may improve bioinformatic and phylogenetic analyses.
 
(a) Map of the study area, including prehistoric fossil sites Estancia Nahuel Huapi (ENH) and Cueva Traful (CT). The geographic distribution of C. sociabilis is shown in light grey. Numbered dots correspond to modern sampling localities: (1) Cerro Monte Redondo; (2) Rincon Grande; (3) Altos del Fortin; (4) Paso Coihue; (5) Cerro la Lagunita. (b) Frequency of haplotypes (based on 253 bp) detected at ENH and CT, presented in time-intervals corresponding to radiocarbon dates for stratigraphic levels. Also shown are haplotype frequencies for the nearest modern collection localities. (c) Relative abundance of C. sociabilis and C. haigi at CT over the last 10 000 years by time-interval. Shading denotes an increase in density of Nothofagus forest at 8500 ybp; dotted line denotes volcanic eruption with ash deposit in CT. (d ) Haplotype networks for cytochrome b sequence data from modern and ancient DNA samples. The sizes of the circles indicate the relative abundance of each haplotype; the numbers of tick marks between circles correspond to the number of base pair substitutions that differentiate haplotypes. Colours correspond to those used in (b) to denote haplotype frequencies.  
Article
Understanding how animal populations have evolved in response to palaeoenvironmental conditions is essential for predicting the impact of future environmental change on current biodiversity. Analyses of ancient DNA provide a unique opportunity to track population responses to prehistoric environments. We explored the effects of palaeoenvironmental change on the colonial tuco-tuco (Ctenomys sociabilis), a highly endemic species of Patagonian rodent that is currently listed as threatened by the IUCN. By combining surveys of modern genetic variation from throughout this species' current geographic range with analyses of DNA samples from fossil material dating back to 10,000 ybp, we demonstrate a striking decline in genetic diversity that is concordant with environmental events in the study region. Our results highlight the importance of non-anthropogenic factors in loss of diversity, including reductions in smaller mammals such as rodents.
 
Article
Pigmy elephants inhabited the islands from the Mediterranean region during the Pleistocene period but became extinct in the course of the Holocene. Despite striking distinctive anatomical characteristics related to insularity, some similarities with the lineage of extant Asian elephants have suggested that pigmy elephants could be most probably seen as members of the genus Elephas. Poulakakis et al (2006) have recently challenged this view by recovering a short mtDNA sequence from an 800 000 year old fossil of the Cretan pigmy elephant (Elephas creticus). According to the authors of this study, a deep taxonomic revision of Cretan dwarf elephants would be needed, as the sequence exhibits clear affinities with woolly mammoth haplotypes. However, we point here many aspects that seriously weaken the strength of the ancient DNA evidence reported.
 
SMR (W ) in relation to mass and manipulated natal brood size (open circles: small broods, 2 or 3 chicks; filled circles: large broods, 5 or 6 chicks). Data shown are the averages per individual over the first and second measurement. 
Residual SMR (W, Cs.e.) in relation to sex and manipulated brood size. Statistical analysis was based on raw data, but the averages shown here were calculated as follows: (i) residuals of regressions of SMR on mass were calculated separately for the first and second measurement, (ii) the mean residual was calculated per individual, and (iii) these means were averaged per group. From left to right, nZ6, 8, 13 and 16. 
Results of the general linear mixed model analysis of SMR (W ). (Series indicates first/second measurement (0 and 1, respectively). Sex is a dummy variable (0, female; 1, male). Top panel shows the model with all significant parameters, bottom panel shows the parameters that were not significant when added to the final model. Note that the significance of the effect of sex changes when the brood size!sex interaction is added together with sex-see text for details.) 
Article
Long-term effects of developmental conditions on health, longevity and other fitness components in humans are drawing increasing attention. In evolutionary ecology, such effects are of similar importance because of their role in the trade-off between quantity and quality of offspring. The central role of energy consumption is well documented for some long-term health effects in humans (e.g. obesity), but little is known of the long-term effects of rearing conditions on energy requirements later in life. We manipulated the rearing conditions in zebra finches (Taeniopygia guttata) using brood size manipulation and cross-fostering. It has previously been shown in this species that being reared in a large brood has negative fitness consequences, and that such effects are stronger in daughters than in sons. We show that, independent of mass, standard metabolic rate of 1-year-old birds was higher when they had been reared in a large brood, and this is to our knowledge the first demonstration of such an effect. Furthermore, the brood size effect was stronger in daughters than in sons. This suggests that metabolic efficiency may play a role in mediating the long-term fitness consequences of rearing conditions.
 
Article
The nomadic Romani (gypsy) people are known for their deep-rooted traditions, but most of their history is recorded from external sources. We find evidence for a Romani genetic lineage in England long before their recorded arrival there. The most likely explanations are that either the historical record is wrong, or that early liaisons between Norse and Romani people during their coincident presence in ninth to tenth century Byzantium led to the spread of the haplotype to England.
 
Morphospace occupation for dinosaurs and crurotarsans. (a) Late Triassic (Carnian-Norian) and (b) Early Jurassic (Hettangian-Toarcian). Triassic and Jurassic taxa are mapped onto a single two-dimensional morphospace, based on the first two principal coordinate axes recovered by the PCO analysis of the entire dataset. Dinosaur morphospace slightly increased in the Early Jurassic, whereas crurotarsan morphospace crashed after the TJ extinction. Note that dinosaur and crurotarsan morphospaces do not overlap: this is an artefact of the discrete character set, which has a phylogenetic structure. Importantly, those crurotarsans heavily convergent with dinosaurs (e.g. poposauroids, 'rauisuchids') are intermediate in morphospace between dinosaurs and the bulk of crurotarsans. 
Morphological disparity for crurotarsans, dinosaurs and avemetatarsalians across the Triassic and the Early Jurassic. (a) Crurotarsans and dinosaurs, (b) crurotarsans and their sister clade, Avemetatarsalia, (c) disparity and diversity for dinosaurs. Error bars denote 95% bootstrap confidence intervals. Norian measures in (b) are shown offset to prevent the loss of information beneath overlapping error bars. Diversity count in (c) is a phylogenetic estimate based on total observed fossils plus implied ghost ranges. 
Article
The evolutionary radiation of dinosaurs in the Late Triassic and Early Jurassic was a pivotal event in the Earth's history but is poorly understood, as previous studies have focused on vague driving mechanisms and have not untangled different macroevolutionary components (origination, diversity, abundance and disparity). We calculate the morphological disparity (morphospace occupation) of dinosaurs throughout the Late Triassic and Early Jurassic and present new measures of taxonomic diversity. Crurotarsan archosaurs, the primary dinosaur 'competitors', were significantly more disparate than dinosaurs throughout the Triassic, but underwent a devastating extinction at the Triassic-Jurassic boundary. However, dinosaur disparity showed only a slight non-significant increase after this event, arguing against the hypothesis of ecological release-driven morphospace expansion in the Early Jurassic. Instead, the main jump in dinosaur disparity occurred between the Carnian and Norian stages of the Triassic. Conversely, dinosaur diversity shows a steady increase over this time, and measures of diversification and faunal abundance indicate that the Early Jurassic was a key episode in dinosaur evolution. Thus, different aspects of the dinosaur radiation (diversity, disparity and abundance) were decoupled, and the overall macroevolutionary pattern of the first 50Myr of dinosaur evolution is more complex than often considered.
 
Article
We present niche-based modelling to project the distribution of 845 European plant species for Germany using three different models and three scenarios of climate and land use changes up to 2080. Projected changes suggested large effects over the coming decades, with consequences for the German flora. Even under a moderate scenario (approx. +2.2 degrees C), 15-19% (across models) of the species we studied could be lost locally-averaged from 2,995 grid cells in Germany. Models projected strong spatially varying impacts on the species composition. In particular, the eastern and southwestern parts of Germany were affected by species loss. Scenarios were characterized by an increased number of species occupying small ranges, as evidenced by changes in range-size rarity scores. It is anticipated that species with small ranges will be especially vulnerable to future climate change and other ecological stresses.
 
Top-cited authors
Céline Bellard
  • Center for biodiversity and environment research, University College of London
Tim M Blackburn
  • University College London
Yves Le Conte
  • French National Institute for Agriculture, Food, and Environment (INRAE)
Cédric Alaux
  • French National Institute for Agriculture, Food, and Environment (INRAE)
Daniel T Blumstein
  • University of California, Los Angeles