The Antarctic krill, Euphausia superba, is an abundant and key species found in the Southern Ocean that forms dense, discrete swarms. Despite over three decades of research on Antarctic krill, the genetics of individual swarms is yet to be specifically investigated. In this study, we address the genetic diversity, population structure and demographic history of nine Antarctic krill swarms by sequencing 1173 bases of the gene cytochrome c oxidase subunit I (cox1, COI) from 504 individuals. Both haplotype diversity (h=0.9974-1.0000) and nucleotide diversity (pi=0.010275-0.011537) of Antarctic krill swarm samples was consistently high compared with populations of other species reported in the literature. Analysis of molecular variance did not show any significant genetic structure, thus implying that the sampled swarms do not appear to reflect discrete genetic units. Fu's Fs and Bayesian Skyride analyses provided strong evidence for a large increase in the population size of Antarctic krill, or selection favouring a particular mitochondrial lineage, within the last few 100 000 years (Pleistocene). The swarm-level results presented in this study not only further our understanding of Antarctic krill biology but, because of the economical importance of this species, also provide data to consider for future krill stock management.Heredity advance online publication, 20 January 2010; doi:10.1038/hdy.2009.188.

Neutral nucleotide diversity does not scale with population size as expected, and this "paradox of variation" is especially severe for animal mitochondria. Adaptive selective sweeps are often proposed as a major cause, but a plausible alternative is selection against large numbers of weakly deleterious mutations subject to Hill-Robertson interference. The mitochondrial genealogies of several species of whale lice (Amphipoda: Cyamus) are consistently too short relative to neutral-theory expectations, and they are also distorted in shape (branch-length proportions) and topology (relative sister-clade sizes). This pattern is not easily explained by adaptive sweeps or demographic history, but it can be reproduced in models of interference among forward and back mutations at large numbers of sites on a nonrecombining chromosome. A coalescent simulation algorithm was used to study this model over a wide range of parameter values. The genealogical distortions are all maximized when the selection coefficients are of critical intermediate sizes, such that Muller's ratchet begins to turn. In this regime, linked neutral nucleotide diversity becomes nearly insensitive to N. Mutations of this size dominate the dynamics even if there are also large numbers of more strongly and more weakly selected sites in the genome. A genealogical perspective on Hill-Robertson interference leads directly to a generalized background-selection model in which the effective population size is progressively reduced going back in time from the present.

When researchers first caught a glimpse of the lush carpet of pink tubeworms covering the scattered bones of a dead grey whale 2900 m below the surface of Monterey Bay, the excitement onboard the Western Flyer (the mother ship of the remotely operated vehicle the Tiburon) must have been electrifying. The discovery of a new genus and several species of whale bone-eating Osedax tubeworms (Annelida, Siboglinidae) a mere 6 years ago from the deep sea was itself noteworthy. But what the researchers peering into the video monitors aboard the Western Flyer could not have known at that moment was that in the gelatinous tubes of those worms clung even more peculiar forms: harems of tiny, paedomorphic males of Osedax, numbering in the hundreds at times. Whereas female tubeworms bore into the marrow of whale bones (possibly via enzymes from their endosymbiotic bacteria), the dwarf males secondarily colonize the tubes of the resident females. The number of males in a female's tube increases over time in a curvilinear fashion. Dwarf males are known from all Osedax species examined to date, yet the origin of the males was an open question. In this issue, Vrijenhoek et al. provide compelling evidence that dwarf males found in the tubes of female Osedax worms are derived from a common larval pool and are unlikely to be the sons of host females or the progeny of females in the local genetic neighbourhood. This study provides an important foundation for future work on the ecology and evolution of extreme male dwarfism in Osedax and sexual size dimorphism more generally.

Reconstructing the history of exploited populations of whales requires fitting a trajectory through at least three points in time:(i) prior to exploitation, when abundance is assumed to be at the maximum allowed by environmental carrying capacity;(ii) the point of minimum abundance or 'bottleneck', usually near the time of protection or the abandonment of the hunt; and (iii) near the present, when protected populations are assumed to have undergone some recovery. As historical abundance is usually unknown, this trajectory must be extrapolated according to a population dynamic model using catch records, an assumed rate of increase and an estimate of current abundance, all of which have received considerable attention by the International Whaling Commission (IWC). Relatively little attention has been given to estimating minimum abundance (N(min)), although it is clear that genetic and demographic forces at this point are critical to the potential for recovery or extinction of a local population. We present a general analytical framework to improve estimates of N(min) using the number of mtDNA haplotypes (maternal lineages) surviving in a contemporary population of whales or other exploited species. We demonstrate the informative potential of this parameter as an a posteriori constraint on Bayesian logistic population dynamic models based on the IWC Comprehensive Assessment of the intensively exploited southern right whales (Eubalaena australis) and published surveys of mtDNA diversity for this species. Estimated historical trajectories from all demographic scenarios suggested a substantial loss of mtDNA haplotype richness as a result of 19th century commercial whaling and 20th century illegal whaling by the Soviet Union. However, the relatively high rates of population increase used by the IWC assessment predicted a bottleneck that was implausibly narrow (median, 67 mature females), given our corrected estimates of N(min). Further, high levels of remnant sequence diversity (theta) suggested that pre-exploitation abundance was larger than predicted by the logistic model given the catch record, which is known to be incomplete. Our results point to a need to better integrate evolutionary processes into population dynamic models to account for uncertainty in catch records, the influence of maternal fidelity on metapopulation dynamics, and the potential for inverse density dependence (an 'Allee effect') in severely depleted populations.

Since its immigration in the Pacific island of New Caledonia in 1942 (i.e. about 240 tick-generations ago), the cattle tick Boophilus microplus has experienced a remarkable adaptive diversification there. In order to better understand the population factors involved, we have investigated the B. microplus population structure on that main host-species, Bos taurus. This study was based microsatellite loci and confirmed that the island colonization came along with a significant bottleneck. Knowledge on B. microplus biology led us to expect B. microplus populations to be composed of highly inbred lineages irregularly dispatched among the individual hosts belonging to the same herds. Instead, this study evidenced a weak inbreeding level and an absence of genetic differentiation within herds. Complementarily, a significant signal of isolation by distance exhibited that human-traffic of cattle does not promote high tick dispersal within the island. Finally, the tick density was found to be about a few hundreds of reproducing adults per squared kilometre, for a gene dispersal range of about a few hundred metres per tick generation. Results are discussed with regard to the evolution of new adaptive changes.

Neutral nucleotide diversity does not scale with population size as expected, and this "paradox of variation" is especially severe for animal mitochondria. Adaptive selective sweeps are often proposed as a major cause, but a plausible alternative is selection against large numbers of weakly deleterious mutations subject to Hill-Robertson interference. The mitochondrial genealogies of several species of whale lice (Amphipoda: Cyamus) are consistently too short relative to neutral-theory expectations, and they are also distorted in shape (branch-length proportions) and topology (relative sister-clade sizes). This pattern is not easily explained by adaptive sweeps or demographic history, but it can be reproduced in models of interference among forward and back mutations at large numbers of sites on a nonrecombining chromosome. A coalescent simulation algorithm was used to study this model over a wide range of parameter values. The genealogical distortions are all maximized when the selection coefficients are of critical intermediate sizes, such that Muller's ratchet begins to turn. In this regime, linked neutral nucleotide diversity becomes nearly insensitive to N. Mutations of this size dominate the dynamics even if there are also large numbers of more strongly and more weakly selected sites in the genome. A genealogical perspective on Hill-Robertson interference leads directly to a generalized background-selection model in which the effective population size is progressively reduced going back in time from the present.

Ocean warming will undoubtedly affect the migratory patterns of many marine species, but specific changes can be predicted only where behavioural mechanisms guiding migration are understood. Southern right whales show maternally inherited site fidelity to near-shore winter nursery grounds, but exactly where they feed in summer (collectively and individually) remains mysterious. They consume huge quantities of copepods and krill, and their reproductive rates respond to fluctuations in krill abundance linked to El Niño Southern Oscillation (ENSO). Here we show that genetic and isotopic signatures, analysed together, indicate maternally directed site fidelity to diverse summer feeding grounds for female right whales calving at Península Valdés, Argentina. Isotopic values from 131 skin samples span a broad range (-23.1 to -17.2‰ δ¹³C, 6.0 to 13.8‰ δ¹⁵N) and are more similar than expected among individuals sharing the same mitochondrial haplotype. This pattern indicates that calves learn summer feeding locations from their mothers, and that the timescale of culturally inherited site fidelity to feeding grounds is at least several generations. Such conservatism would be expected to limit the exploration of new feeding opportunities, and may explain why this population shows increased rates of reproductive failure in years following elevated sea-surface temperature anomalies off South Georgia, the richest known feeding ground for baleen whales in the South Atlantic.

The population structure and mitochondrial (mt) DNA diversity of southern right whales (Eubalaena australis) are described from 146 individuals sampled on 4 winter calving grounds (Argentina, South Africa, Western Australia, and the New Zealand sub-Antarctic) and 2 summer feeding grounds (South Georgia and south of Western Australia). Based on a consensus region of 275 base pairs of the mtDNA control region, 37 variable sites defined 37 unique haplotypes, of which only one was shared between regional samples of the Indo-Pacific and South Atlantic Oceans. Phylogenetic reconstruction of the southern right whale haplotypes revealed 2 distinct clades that differed significantly in frequencies between oceans. An analysis of molecular variance confirmed significant overall differentiation among the 4 calving grounds at both the haplotype and the nucleotype levels (F(ST)= 0.159; Phi(ST)= 0.238; P < 0.001). Haplotype diversity was significantly lower in the Indo-Pacific (h = 0.701 +/- 0.037) compared with the South Atlantic (h = 0.948 +/- 0.013), despite a longer history of exploitation and larger catches in the South Atlantic. In fact, the haplotype diversity in the Indo-Pacific basin was similar to that of the North Atlantic right whale that currently numbers about 300 animals. Multidimensional scaling of genetic differentiation suggests that gene flow occurred primarily between adjacent calving grounds within an ocean basin, with mixing of lineages from different calving grounds occurring on feeding grounds.

Vessel-whale collision events represented the ultimate cause of death for 21 (52.5%) of the 40 North Atlantic right whales (Eubalaena glacialis) necropsied between 1970 and December 2006. Injuries seen in vessel-struck whales fall into two distinct categories: 1) sharp trauma, often resulting from contact with the propeller, and 2) blunt trauma, presumably resulting from contact with a vessel's hull. This study analyzes four trauma cases that resulted from vessel-whale collisions, which together provide a framework for a more critical understanding of lethal blunt and sharp trauma resulting from vessel collisions with right whales. In case no. 1, contact with a propeller resulted in three deep lacerations. The animal survived acute trauma only to succumb nearly 14 years later when the lesions reopened and became infected. In case no. 2, anecdotal reports linked the laceration of large arteries of the peduncle and histologic evidence of perimortem trauma at a bone fracture site to vessel-whale collision trauma. Case no. 3 had a laceration of the oral rete and a fracture of the rostrum. Both of the areas displayed histologic evidence of perimortem blunt trauma. Finally, in case no. 4, an antemortem mandibular fracture, two additional skull fractures, and widespread hemorrhage were consistent with severe blunt trauma. Evidence from each case, including the timing of trauma relative to the time of death and identifying characteristics of both trauma types, are presented. Before this study, no detailed comparative analysis of trauma pathology that resulted from lethal interactions between vessels and right whales had been conducted. This study demonstrates the importance of detailed gross and histologic examination in determining the significance and timing of traumatic events. This work represents a new paradigm for the differential diagnosis of lethal sharp and blunt trauma in right whales hit by ships and will enhance the present understanding of the impact of anthropogenic mortality on this critically endangered species.

  United States and Canadian governments have responded to legal requirements to reduce human-induced whale mortality via vessel strikes and entanglement in fishing gear by implementing a suite of regulatory actions. We analyzed the spatial and temporal patterns of mortality of large whales in the Northwest Atlantic (23.5°N to 48.0°N), 1970 through 2009, in the context of management changes. We used a multinomial logistic model fitted by maximum likelihood to detect trends in cause-specific mortalities with time. We compared the number of human-caused mortalities with U.S. federally established levels of potential biological removal (i.e., species-specific sustainable human-caused mortality). From 1970 through 2009, 1762 mortalities (all known) and serious injuries (likely fatal) involved 8 species of large whales. We determined cause of death for 43% of all mortalities; of those, 67%(502) resulted from human interactions. Entanglement in fishing gear was the primary cause of death across all species (n= 323), followed by natural causes (n= 248) and vessel strikes (n= 171). Established sustainable levels of mortality were consistently exceeded in 2 species by up to 650%. Probabilities of entanglement and vessel-strike mortality increased significantly from 1990 through 2009. There was no significant change in the local intensity of all or vessel-strike mortalities before and after 2003, the year after which numerous mitigation efforts were enacted. So far, regulatory efforts have not reduced the lethal effects of human activities to large whales on a population-range basis, although we do not exclude the possibility of success of targeted measures for specific local habitats that were not within the resolution of our analyses. It is unclear how shortfalls in management design or compliance relate to our findings. Analyses such as the one we conducted are crucial in critically evaluating wildlife-management decisions. The results of these analyses can provide managers with direction for modifying regulated measures and can be applied globally to mortality-driven conservation issues.

Molting in arthropods is orchestrated by a series of endocrine changes that occur towards the end of an instar. However, little is understood about the mechanisms that trigger these endocrine changes. Here, nutritional inputs were manipulated to investigate the minimal nutritional inputs required for a Manduca sexta larva to initiate a molt. Amino acids were found to be necessary for a larva to molt, indicating the involvement of an amino acid sensitive pathway. Feeding rapamycin, an inhibitor of the target of rapamycin (TOR) signaling, delayed the onset of a molt and resulted in abnormally larger larvae. Rapamycin also suppressed the growth of the prothoracic glands relative to the whole body growth, and this was accompanied by suppression of ecdysone production and secretion. Higher doses of rapamycin also slowed the growth rate, indicating that TOR signaling also plays a role in systemic growth. TOR signaling therefore couples the nutritional status of the larva to the endocrine system to regulate the timing of a molt.

The traditional taxonomy of ca. 700 cone snails assigns all species to a single genus, Conus Linnaeus 1758. However, an increasing body of evidence suggests that some belong to a phylogenetically distinct clade that is sometimes referred to as Conasprella. Previous work (Kraus et al. 2011) showed that a short (259 bp) conserved intronic sequence (CIS) of the γ-glutamyl carboxylase gene (intron 9) can be used to delineate deep phylogenetic relationships among some groups of Conus. The work described here uses intron 9 (338 bp) to resolve problematic relationships among the conasprellans and to distinguish them from Conus proper. Synapomorphic mutations at just 39 sites can resolve several groups within Conasprella because the informative region of intron 9 is so well conserved that the phylogenetic signal is not obscured by homoplasies at conflicting sites. Intron 9 also unambiguously distinguishes Conasprella as a whole from Conus because the conserved regions that are so well conserved within each group are not alignable and clearly not homologous between them. This pattern suggests that expression of the γ-glutamyl carboxylase gene may have undergone a functionally significant change in Conus or Conasprella shortly after they diverged.

This paper presents a qualitative investigation of peer mentoring among HIV seropositive injection drug users in a randomized controlled trial, the INSPIRE study. Qualitative analyses of 68 in-depth open-ended interviews conducted in 2005 in Baltimore, New York, Miami, and San Francisco revealed that these individuals conceptualized themselves as change agents through the identity of peer mentor at the three related domains of individual, interpersonal, and community-level change. Implications for program development and future research of peer mentoring as a mechanism for HIV prevention are discussed. This study was funded by the Centers for Disease Control and Prevention and Health Resources and Services Administration (HRSA).

Mitochondrial genes in animals are especially useful as molecular markers for the reconstruction of phylogenies among closely related taxa, due to the generally high substitution rates. Several insect orders, notably Hymenoptera and Phthiraptera, show exceptionally high rates of mitochondrial molecular evolution, which has been attributed to the parasitic lifestyle of current or ancestral members of these taxa. Parasitism has been hypothesized to entail frequent population bottlenecks that increase rates of molecular evolution by reducing the efficiency of purifying selection. This effect should result in elevated substitution rates of both nuclear and mitochondrial genes, but to date no extensive comparative study has tested this hypothesis in insects. Here we report the mitochondrial genome of a crabronid wasp, the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae), and we use it to compare evolutionary rates among the four largest holometabolous insect orders (Coleoptera, Diptera, Hymenoptera, Lepidoptera) based on phylogenies reconstructed with whole mitochondrial genomes as well as four single-copy nuclear genes (18S rRNA, arginine kinase, wingless, phosphoenolpyruvate carboxykinase). The mt-genome of P. triangulum is 16,029 bp in size with a mean A+T content of 83.6%, and it encodes the 37 genes typically found in arthropod mt genomes (13 protein-coding, 22 tRNA, and two rRNA genes). Five translocations of tRNA genes were discovered relative to the putative ancestral genome arrangement in insects, and the unusual start codon TTG was predicted for cox2. Phylogenetic analyses revealed significantly longer branches leading to the apocritan Hymenoptera as well as the Orussoidea, to a lesser extent the Cephoidea, and, possibly, the Tenthredinoidea than any of the other holometabolous insect orders for all mitochondrial but none of the four nuclear genes tested. Thus, our results suggest that the ancestral parasitic lifestyle of Apocrita is unlikely to be the major cause for the elevated substitution rates observed in hymenopteran mitochondrial genomes.

The International Whaling Commission (IWC) is currently engaged in an intensive negotiating process in an attempt to resolve international disputes about whaling. The IWC has pioneered and agreed a management procedure approach for setting catch limits for commercial whaling that was unanimously recommended by its Scientific Committee. It is disturbing that current negotiations are moving towards discarding this agreed and carefully developed scientific procedure in favour of ad hoc catch allowances based on political expediency.

Making a life-sized model of a blue whale is difficult. Making one to the exacting standards of 'scientific accuracy' is backbreaking. When, in the early 1960s, the American Museum of Natural History in New York undertook to fabricate a replica of the largest animal that ever lived, little was known about how blue whales really looked and behaved in the wild. Exhibitors like Richard Van Gelder guided themselves by old photographs, illustrations, tables of measurements and the experience of other institutions--as well as their own educated guesswork, and ideas of beauty, value and pride.

Analysis of one year of acoustic signal recordings from the five permanent autonomous hydrophones of the International Monitoring System in the South-West Indian Ocean reveals low frequency with high intensity calls produced by two blue whale subspecies. The "Antarctic" or "true" blue whale (B. m. intermedia) calls and the "Madagascar-type" Pygmy blue whale calls (B. m. brevicauda) were automatically detected through the matched filtering method. The potential movements were investigated by using the time difference of arrival (TDOA) of calls to assess the bearing of the sound source. The fully range dependent parabolic equation code (RAM - range-dependent acoustic model) and the PMCC code (progressive multi-channel correlation) are applied to estimate the range between our system and the vocalising animals. Our results show that (1) the variation of call number revealed two distinct patterns of seasonal whale occurrences and (2) the distances from the hydrophones to the blue whales reached up to 50 km. Tracking whales is possible when whales are concentrated of the hydrophone array.

Understanding the group behavior of free-swimming whales is critical to the management and conservation of many whale species. Moreover, understanding how whales coordinate their activities could serve as a model for improving the communication and navigation systems of underwater robots and autonomous underwater vehicles (AUVs). One method for studying the group behavior of free-swimming whales involves simultaneously attaching acoustic recording tags to multiple animals within a close-swimming conspecific group. Acoustic recording tags can sample an animal's acoustic environment simultaneously with other physical parameters such as diving depth and acceleration. In this paper, methods are presented for dynamically tracking whales that vocalize regularly such as odontocetes using the DTAG, a digital acoustic recording tag developed by the Woods Hole Oceanographic Institution. A trilateration model is reviewed for tracking groups of whales that include three or more tagged whales using time of flight data and results are given from applying this model to a novel data set of three simultaneously diving and cross-audible short-finned pilot whales (Globicephala macrorhyncus). A new model is subsequently proposed for tracking whale groups that include only two tagged whales using a generalization of multilateration and data captured by stereo DTAGs.

Eastern Pacific grey whales (Eschrichtius robustus) apparently do not actively echolocate, yet still they thrive in shallow water environments where visibility is much reduced. Along their migration route and in their feeding grounds these whales are exposed to high levels of ambient noise, highlyturbid waters and many underwater obstacles. To test possible passive acoustic localization mechanisms (e.g. Acoustic Daylight Imaging and Passive Synthetic Aperture), we made extensive acoustic measurements during comprehensive field studies of these whales on their summer feeding grounds in British Columbia (Canada). In combination with visual observations of the whales and their behaviours, we investigated the acoustical sources available to the whales when navigating within a feeding bay. First, we measured ambient noise levels to construct the acoustic landscape around the whales. Second, we investigated how sound is altered when objects such as kelp beds and rocks are present. We also measured acoustic changes induced by direct, controlled modifications of the near-shore environment. The aim of this research is to understand how grey whales might be finding their way around, and what impact, if any, increased levels of ambient noise might have on the whales' ability to find food and navigate within the feeding grounds.

We report initial results from a study on behavioral responses of beaked and other whales to sonar and other sounds. This research is designed to provide new science-based approaches for mitigating the risk of sonar to beaked and other whales. The study was conducted at the Atlantic Undersea Test and Evaluation Center (AUTEC) range near Andros Island in the Bahamas, where Blainville's beaked whales (Mesoplodon densirostris) can regularly be detected using passive acoustic monitoring of their echolocation clicks. Tags recorded sound at the whale and behavior of the whale. Data were collected from 10 tag deployments, 6 on Blainville's beaked whales and 4 on pilot whales. 109 hours of data were collected from tags, 74 hours from beaked whales and 34 hours from pilot whales. Playbacks of mid-frequency sonar and killer whale calls were performed on 3 of the tagged whales, 1 beaked whale and 2 pilot whales. The tagged beaked whale responded to both sonar and killer whale sounds by premature cessation of clicking during foraging dives (RL = approximately 117 dB re 1 muPa for the killer whale sound, approximately 145 dB for the sonar), and an unusually slow and long ascent.

The use of voluntary approaches to achieve conservation goals is becoming increasingly popular. Nevertheless, few researchers have quantitatively evaluated their efficacy. In 1998 industry, government agencies, and nongovernmental organizations established a voluntary conservation program for whale watching in the northeast region of the United States, with the intent to avoid collisions with and harassment of endangered whales by commercial and recreational whale-watching vessels. One important aspect of the program was the establishment of 3 speed zones within specific distances of whales. We wanted to determine the level of compliance with this aspect of the program to gauge its efficacy and gain insights into the effectiveness of voluntary measures as a conservation tool. Inconspicuous observers accompanied 46 commercial whale-watching trips from 12 companies in 2003 (n= 35) and 2004 (n= 11). During each trip, vessel position and speed were collected at 5-second intervals with a GPS receiver. Binoculars with internal laser rangefinders and digital compasses were used to record range and bearing to sighted whales. We mapped whale locations with ArcGIS. We created speed-zone buffers around sighted whales and overlaid them with vessel-track and speed data to evaluate compliance. Speeds in excess of those recommended by the program were considered noncompliant. We judged the magnitude of noncompliance by comparing a vessel's maximum speed within a zone to its maximum recorded trip speed. The level of noncompliance was high (mean 0.78; company range 0.74-0.88), some companies were more compliant than others (p= 0.02), noncompliance was significantly higher in zones farther from whales (p < 0.001), and operators approached the maximum speed capabilities of their vessel in all zones. The voluntary conservation program did not achieve the goal of substantially limiting vessel speed near whales. Our results support the need for conservation programs to have quantifiable metrics and frequent evaluation to ensure efficacy.

The strength of selection to increase the span of a life stage is dependent upon individuals at that stage being able to contribute towards individual fitness and the probability of their surviving to that stage. Complete reproductive cessation and a long post-reproductive female lifespan as found in humans are also found in killer whale (Orcinus orca) and short-finned pilot whale (Globicephala macrorhynchus), but not in the long-finned pilot whale (Globicephala melaena). Each species forms kin-based, stable matrilineal groups and exhibits kin-directed behaviours that could increase inclusive fitness. Here, the initial mortality rate and mortality rate-doubling time of females of these three closely related whale species are compared. The initial mortality rate shows little variation among pilot whale species; however mortality rate accelerates almost twice as fast in the long-finned pilot whale as it does in killer whale and short-finned pilot whale. Selection for a long post-reproductive female lifespan in matrilineal whales may therefore be determined by the proportion of females surviving past the point of reproductive cessation.