Gastric evacuation in the sandbar shark, Carcharhinus plumbeus

Sandbar sharks maintained in an enclosure in the natural environment were fed meals of soft blue crab, Callinectes sapidus , or menhaden, Brevoortia lyrannus , and were sacrificed at intervals after feeding to measure rates of gastric emptying. Regression analysis was used to evaluate the adequacy of various models in describing the decrease in stomach contents with time after feeding. A Gompertz growth curve provided the best fit to the data for both food types. This model suggests that gastric emptying is characterized by an initial lag phase during which evacuation rate increases, a period of maximal evacuation, and a decreasing evacuation rate during the later stages of digestion. Passage of food from the stomach was slow relative to other species of fish studied. The time required to evacuate 98% of a meal was 92–3 and 70–7 h for sharks fed menhaden and blue crab respectively. Evacuation rates did not differ between day and night periods.     

Histological and morphological aspects of reproduction in the sandbar shark Carcharhinus plumbeus in the U.S. south‐eastern Atlantic Ocean and Gulf of Mexico

The reproduction of the sandbar shark Carcharhinus plumbeus in the U.S. south‐eastern Atlantic Ocean including the Gulf of Mexico was examined using a combination of histological and morphological characteristics of C. plumbeus collected through fishery‐dependent and ‐independent sampling programmes (n = 1,567). Indices of maturity were constructed using measurements of gonads, reproductive tracts and claspers, and sandbar sharks exhibited 50% maturity sizes of 140 and 148 cm fork length for males and females respectively. Gonado‐somatic indices and variation in reproductive tract condition were used to determine seasonal trends in reproduction of mature C. plumbeus. Sandbar sharks have discrete seasonal reproductive cycles in which males produce sperm from January to May with a peak in May and females develop eggs from January to May with ovulation occurring in June. Females were shown to exhibit a >2 year reproductive cycle. Embryonic development was assessed through measurements of masses and lengths of uterine contents. Gestation was 12 months, from July to the following June, with parturition in late June. This research highlights a difference from previously reported data on the periodicity of female reproduction in C. plumbeus in the U.S. south‐eastern Atlantic Ocean and Gulf of Mexico, which may have major effects on future C. plumbeus stock management.     

Teeth penetration force of the tiger shark Galeocerdo cuvier and sandbar shark Carcharhinus plumbeus

This study examined the minimum force required of functional teeth and replacement teeth in the tiger shark Galeocerdo cuvier and the sandbar shark Carcharhinus plumbeus to penetrate the scales and muscle of sheepshead Archosargus probatocephalus and pigfish Orthopristis chrysoptera. Penetration force ranged from 7·7–41·9 and 3·2–26·3 N to penetrate A. probatocephalus and O. chrysoptera, respectively. Replacement teeth required significantly less force to penetrate O. chrysoptera for both shark species, most probably due to microscopic wear of the tooth surfaces supporting the theory shark teeth are replaced regularly to ensure sharp teeth that are efficient for prey capture.     

Isolation and characterization of five dinucleotide microsatellite loci in the sandbar shark,Carcharhinus plumbeus

Five dinucleotide markers were isolated and optimized from a microsatellite‐enriched genomic library obtained from the sandbar shark, Carcharhinus plumbeus. Genotypic distributions of all markers were found to be in conformance with the expectations of Hardy–Weinberg equilibrium with four to 39 alleles present per locus. We amplified these loci in two female sharks and their litters. A maternal allele was recovered at each locus in all progeny indicating reliable amplification. More than two paternal alleles were recovered across both litters indicating genetic polyandry. Additionally, these markers were amplified across 10 carcharhiniform species to examine their utility in other studies.     

Global phylogeography of the scalloped hammerhead shark (Sphyrna lewini)

Large marine fishes typically have little population genetic structure. The exceptions are associated with sedentary behaviour, disjunct distributions, or reproductive philopatry. Scalloped hammerhead sharks (Sphyrna lewini) incorporate the contrasting traits of oceanic habitat (usually associated with high dispersal) and possible fidelity to nursery grounds (for reproductive females). To evaluate the expectations of these contrasting behaviours, we examined the global genetic structure of S. lewini based on collections (n = 271 individuals) from 20 nursery areas. A 548-bp fragment of mitochondrial DNA control region revealed 22 polymorphic sites, 24 haplotypes, and three lineages distinguished by 2.56-3.77% sequence divergence. Coalescence analyses based on a provisional molecular clock indicate an origin in the Indo-West Pacific with late Pleistocene radiations into the central Pacific (Hawaii) and eastern Pacific (Central America), as well as recent interchange between oceans via southern Africa. Population subdivisions are strong (overall Phi(ST) = 0.749, P < 0.0001 and among oceans Phi(ST) = 0.598, P < 0.0098). Genetic discontinuity within oceans (Phi(ST) = 0.519, P < 0.0001) is primarily associated with oceanic barriers (migration across oceans M approximately 0), with much less structure along continental margins (M > 10). We conclude that nursery populations linked by continuous coastline have high connectivity, but that oceanic dispersal by females is rare. Although we cannot rule out philopatry to natal nurseries, oceanic barriers appear to have a much stronger influence on the genetic architecture of this species and may indicate a mechanism for recent evolutionary radiations in the genus Sphyrna.     

Food habits and ontogenetic changes in the diet of the sandbar shark, Carcharhinus plumbeus, in Hawaii

The sandbar shark, Carcharhinus plumbeus, is a wide-ranging coastal species in tropical and temperate regions, and it is the most common species of shark in Hawaii, as in many locations where it occurs. Information on the diet and feeding habits of this species in the Pacific Ocean are extremely limited. For this study we quantified the diet of sandbar sharks in Hawaii based on records collected during the Hawaii Cooperative Shark Research and Control Program from 1967 to 1969. During this program a total of 565 stomachs were examined, of which 265 contained food. Sharks ranged in size from 59 to 190 cm total length. Teleosts were the most common prey group, but both cephalopods and crustaceans also occurred frequently. Ontogenetic changes in diet of sandbar sharks were apparent, with crustaceans forming a greater proportion of the diet of smaller sharks. Both cephalopods and elasmobranchs increased in importance with increasing shark size. Prey diversity also increased with size, with large, mobile, and reef prey species found more commonly in the diet of larger sharks. Mature male and female sharks appeared to segregate by depth, though major differences in the diet between the sexes were not apparent. However, there was some evidence of dietary differences between sharks caught in different depths and seasons. The results of this study suggest that sandbar sharks in Hawaii and throughout the world, are primarily piscivores, but also consume a variety of invertebrate prey, and that their diet varies with geographical location and stage of development.     

Age and growth of the sandbar shark, Carcharhinus plumbeus, in Hawaiian waters through vertebral analysis

Age and growth estimates were determined for the sandbar shark, Carcharhinus plumbeus, from Oahu, Hawaii in the central Pacific Ocean. Age estimates were obtained through vertebral centra analysis of 187 sharks. We verified our age estimates through marginal increment analysis of centra and oxytetracycline marking methods of at liberty sandbar sharks. Sizes of sampled sharks ranged from 46 to 147 cm pre-caudal length. Four growth models were fitted to length-at-age data; two forms of the von Bertalanffy growth model, the Gompertz growth model, and a logistic growth model. Males and females exhibited statistically significant differences in growth, indicating that females grow slower and attain larger sizes than males. Growth parameter estimates revealed slower growth rates than previously estimated (based on captive specimens) for Hawaiian sandbar sharks. The von Bertalanffy growth model using empirical length-at-birth provided the best biological and statistical fit to the data. This model gave parameter estimates of L _∞ = 138.5 cm PCL and k = 0.12 year⁻¹ for males and L _∞ = 152.8 cm PCL, k = 0.10 year⁻¹ for females. Male and female sandbar sharks mature at approximately 8 and 10 years of age, respectively.     

Genetic polyandry and sexual conflict in the sandbar shark, Carcharhinus plumbeus, in the western North Atlantic and Gulf of Mexico

To investigate patterns of polyandry in the sandbar shark (Carcharhinus plumbeus), 20 pregnant females were sampled from the western North Atlantic and Gulf of Mexico. Five species-specific microsatellite markers were used to genotype each shark and its litter. Of 20 litters, 17 (85%) were shown to have multiple sires. In multiply sired litters, the estimated minimum number of sires ranged from two to five with an average of 2.3 males per litter. Regression analysis did not demonstrate a significant relationship between female reproductive success and female body size or sire number and female body size. There was a high incidence of reproductive skew noted in litters, and two groups of males with significantly different mean reproductive success were observed. Analyses using Bateman's principles suggest that there is less direct benefit for females that acquire multiple mates than for males who bias paternity within litters. In light of past morphological and behavioural studies, these data suggest that patterns of polyandry in elasmobranchs may be determined by coercive mating, and that breeding behaviour has likely evolved in the context of sexual conflict.     

Effective size closely approximates the census size in the heavily exploited western Atlantic population of the sandbar shark, Carcharhinus plumbeus

The sandbar shark, Carcharhinus plumbeus, is a long-lived species with low lifetime fecundity that is heavily fished in the western North Atlantic. Inshore nursery grounds increase survivorship of sandbar shark pups and the principal nurseries are in the mid-Atlantic region. We calculated effective number of breeders (N_b) and effective population size (N_e) for adults utilizing the nursery grounds of the Delaware Bay and the Eastern Shore of Virginia by genotyping 902 animals across five cohorts at eight microsatellite loci. Estimates of N_b and N_e were compared to estimates of census size (N_c) of cohorts obtained from Delaware Bay. The estimated N_e/N_c and N_b/N_c ratios were 0.45 or higher whether the Delaware Bay cohorts were considered as distinct year classes or combined. This is in contrast to estimated N_e/N_c ratios in other exploited marine fishes, which are several orders of magnitude smaller. Instead, the N_e/N_c ratio of sandbar sharks is similar to that found in marine and terrestrial mammals.     

A role for nonphysical barriers to gene flow in the diversification of a highly vagile seabird, the masked booby (Sula dactylatra)

To test the hypothesis that nonphysical barriers to gene flow play a role in the divergence of low-latitude seabird populations, we applied phylogeographic methods to mitochondrial control region sequence variation in a global sample of masked boobies (Sula dactylatra). In accord with previous studies, we found that Indo-Pacific and Atlantic haplotypes form two divergent lineages, excluding one haplotype previously attributed to secondary contact between the Indian Ocean and the Caribbean Sea. Within the Indo-Pacific and the Atlantic, we found a relatively large number of haplotypes, many of which were unique to a single population. Although haplotypes from most populations were found in more than one higher-level clade, nested clade analysis revealed a significant association between clades and geography for the majority of higher-level clades, most often interpreted as a consequence of isolation by distance. We found low levels of gene flow within Indo-Pacific and Atlantic populations, and a significant correlation between gene flow and geographical distance among Indo-Pacific populations. We estimate that Indo-Pacific masked boobies experienced rapid population growth approximately 180,000 years ago and that the majority of Indo-Pacific and Atlantic populations diverged within the last approximately 115,000 years. These combined data suggest that the predominant pattern between Indo-Pacific and Atlantic populations is long-term isolation by physical barriers to gene flow. In contrast, populations within these regions appear to have diverged despite few obvious physical barriers to gene flow, perhaps as a consequence of limited natal dispersal combined with local adaptation and/or genetic drift.     

Modelling pollen‐mediated gene flow in rice: risk assessment and management of transgene escape

Fast development and commercialization of genetically modified plants have aroused concerns of transgene escape and its environmental consequences. A model that can effectively predict pollen‐mediated gene flow (PMGF) is essential for assessing and managing risks from transgene escape. A pollen‐trap method was used to measure the wind‐borne pollen dispersal in cultivated rice and common wild rice, and effects of relative humidity, temperature and wind speed on pollen dispersal were estimated. A PMGF model was constructed based on the pollen dispersal pattern in rice, taking outcrossing rates of recipients and cross‐compatibility between rice and its wild relatives into consideration. Published rice gene flow data were used to validate the model. Pollen density decreased in a simple exponential pattern with distances to the rice field. High relative humidity reduced pollen dispersal distances. Model simulation showed an increased PMGF frequency with the increase of pollen source size (the area of a rice field), but this effect levelled off with a large pollen‐source size. Cross‐compatibility is essential when modelling PMGF from rice to its wild relatives. The model fits the data well, including PMGF from rice to its wild relatives. Therefore, it can be used to predict PMGF in rice under diverse conditions (e.g. different outcrossing rates and cross‐compatibilities), facilitating the determination of isolation distances to minimize transgene escape. The PMGF model may be extended to other wind‐pollinated plant species such as wheat and barley.     

Huffmanela cf. carcharhini (Nematoda: Trichosomoididae: Huffmanelinae) from skin of a sandbar shark, Carcharhinus plumbeus, in the Pacific Ocean

Eggs of Huffmanela cf. carcharhini from the skin of an aquarium-held, juvenile sandbar shark, Carcharhinus plumbeus , from the Pacific Ocean were studied using light and scanning electron microscopy. Grossly, eggs imparted a scribble-like skin marking approximately 130 × 60 mm on the right side of the shark's snout adjacent to its eye and nostril. Fresh (unfixed) eggs were elliptical, 75-95 μm long (xˉ = 85 μm, SD = ±4.5; n = 75), 48-63 μm wide (53 ± 3.4; 75), 8-10 μm in shell thickness (9 ± 1.3; 27), 45-68 μm in vitelline mass length (52 ± 6.9; 8); had a smooth shell surface and nonprotruding polar plugs 8-13 μm wide (10 ± 1.5; 73); lacked thin filaments, superficial envelope, and shell spines; sank in 35 ppt artificial seawater; and did not spontaneously hatch after 12 hr in 35 ppt artificial seawater. Formalin-fixed eggs measured 193 days postfixation were 75-95 μm long (84 ± 3.9; 150), 45-60 μm wide (50 ± 2.2; 150), 5-10 μm in shell thickness (8 ± 1.2; 87), 45-60 μm in vitelline mass length (51 ± 3.0; 92), and 30-40 μm in vitelline mass width (33 ± 2.0; 84), and had nonprotruding polar plugs that were 10-15 μm long (11 ± 1.4; 93) and 8-10 μm wide (9 ± 1.1; 108). Forcibly hatched first-stage larvae (unfixed) were filiform, 188-273 μm long (212 ± 25.5; 13), 8-13 μm wide (10 ± 1.2; 13), and had fine transverse striations. Eggs infected the epidermis only. Histology revealed intra-epithelial inflammation with eosinophilic granulocytes and hyperplasia, plus dermal lymphofollicular hyperplasia associated with the infection. The eggs of H. cf. carcharhini likely undergo considerable ex utero development before being sloughed (unhatched) from the host, along with epidermal cells.     

Historical male‐mediated introgression in horseshoe bats revealed by multilocus DNA sequence data

Instances of hybridization between mammalian taxa in the wild are rarely documented. To test for introgression between sibling species of horseshoe bat (Rhinolophus yunanensis and R. pearsoni) and two subspecies of the latter (R. p. pearsoni and R. p. chinensis), we sequenced two mtDNA and two ncDNA markers in individuals sampled from multiple localities within their overlapping ranges. The interspecific mtDNA gene tree corresponded to the expected taxonomic divisions, and coalescent‐based analyses suggested divergence occurred around 4 MYA. However, these relationships strongly conflicted with those recovered from two independent nuclear gene trees, in which R. yunanensis clustered with R. p. pearsoni to the exclusion of R. p. chinensis. This geographically widespread discordance is best explained by large‐scale historical introgression of ncDNA from R. yunanensis to R. pearsoni by male‐mediated exchange in mixed species colonies during Pleistocene glacial periods, when ranges may have contracted and overlapped more than at present. Further species tree–gene tree conflicts were detected between R. p. pearsoni and R. p. chinensis, also indicating past and/or current introgression in their overlapping regions. However, here the patterns point to asymmetric mtDNA introgression without ncDNA introgression. Analyses of coalescence times indicate this exchange has occurred subsequent to the divergence of these subspecies from their common ancestor. Our work highlights the importance of using multiple data sets for reconstructing phylogeographic histories and resolving taxonomic relationships.     

Evidence for contemporary and historical gene flow between guppy populations in different watersheds,with a test for associations with adaptive traits

In dendritic river systems, gene flow is expected to occur primarily within watersheds. Yet, rare cross‐watershed transfers can also occur, whether mediated by (often historical) geological events or (often contemporary) human activities. We explored these events and their potential evolutionary consequences by analyzing patterns of neutral genetic variation (microsatellites) and adaptive phenotypic variation (male color) in wild guppies (Poecilia reticulata) distributed across two watersheds in northern Trinidad. We found the expected signatures of within‐watershed gene flow; yet we also inferred at least two instances of cross‐watershed gene flow—one in the upstream reaches and one further downstream. The upstream cross‐watershed event appears to be very recent (41 ± 13 years), suggesting dispersal via recent flooding or undocumented human‐mediated transport. The downstream cross‐watershed event appears to be considerably older (577 ± 265 years), suggesting a role for rare geological or climatological events. Alongside these strong signatures of both contemporary and historical gene flow, we found little evidence of impacts on presumably adaptive phenotypic differentiation, except perhaps in the one instance of very recent cross‐watershed gene flow. Selection in this system seems to overpower gene flow—at least on the spatiotemporal scales investigated here.     

Patterns of colonization, evolution and gene flow in species of the genus Patella in the Macaronesian Islands

The study of phylogeographical patterns may contribute to a better understanding of factors affecting the dispersal of organisms in ecological and historical times. For intertidal organisms, islands are particularly suitable models allowing the test of predictions related to the efficacy of pelagic larvae dispersal. Here, we study the phylogeographical patterns and gene flow within three groups of species of the genus Patella present in the Macaronesian Islands that have been previously shown to be monophyletic. The genetic variability of around 600 bp of the mitochondrial gene cytochrome c oxidase subunit I was studied by single strand conformation polymorphism and/or sequencing for seven species of limpets. A total of 420 samples were analysed from the Macaronesian archipelagos, North Africa, and Atlantic and Mediterranean shores of the Iberian Peninsula. No clear geographical pattern or temporal congruence was found between the three groups of species, pointing to independent histories and colonization events. However, for the three groups, the split between the Macaronesian and the mainland forms most probably occurred before 3.9 million years ago, predating the establishment of the current circulation patterns. The presence of pelagic larvae in these species is shown to be insufficient to ensure gene flow between continental and Macaronesian populations and between the Macaronesian archipelagos. In the endangered Azorean populations of Patella candei, there is restricted gene flow to Flores and Graciosa.     

Vicariance divergence and gene flow among islet populations of an endemic lizard

Allopatry and allopatric speciation can arise through two different mechanisms: vicariance or colonization through dispersal. Distinguishing between these different allopatric mechanisms is difficult and one of the major challenges in biogeographical research. Here, we address whether allopatric isolation in an endemic island lizard is the result of vicariance or dispersal. We estimated the amount and direction of gene flow during the divergence of isolated islet populations and subspecies of the endemic Skyros wall lizard Podarcis gaigeae, a phenotypically variable species that inhabits a major island and small islets in the Greek archipelago. We applied isolation-with-migration models to estimate population divergence times, population sizes and gene flow between islet-mainland population pairs. Divergence times were significantly correlated with independently estimated geological divergence times. This correlation strongly supports a vicariance scenario where islet populations have sequentially become isolated from the major island. We did not find evidence for significant gene flow within P. g. gaigeae. However, gene-flow estimates from the islet to the mainland populations were positively affected by islet area and negatively by distance between the islet and mainland. We also found evidence for gene flow from one subspecies (P. g. weigandi) into another (P. g. gaigeae), but not in the other direction. Ongoing gene flow between the subspecies suggests that even in this geographically allopatric scenario with the sea posing a strong barrier to dispersal, divergence with some gene flow is still feasible.     

Subdivided population structure and phylogeography of an endangered freshwater snail, Notopala sublineata (Conrad, 1850) (Gastropoda: Viviparidae), in Western Queensland, Australia

Populations of the Australian freshwater snail Notopala sublineata (Conrad, 1850) have declined rapidly over the last decade, but are still abundant in most river systems of Western Queensland. These rivers are characterized by the unpredictable and highly variable nature of their climatic and hydrological regimes, with episodic periods of very large flow and many periods of little or no flow. We used mitochondrial sequences and allozymes to investigate the genetic structure and infer patterns of dispersal of N. sublineata within this unique environment. We sampled 24 waterholes throughout the four major catchments of the Lake Eyre Basin. Based on a 457-bp fragment of the mitochondrial cytochrome oxidase subunit I gene, we identified 55 haplotypes in a sample of 256 individuals. Both nuclear and mitochondrial genetic datasets indicated high levels of genetic subdivision and restricted gene flow among populations within and among catchments. The mitochondrial haplotypes clustered into two main geographical clades, corresponding with two groups of adjacent catchments: Cooper–Bulloo and Diamantina–Georgina, which appear to have diverged 300 000 years ago. Populations of N. sublineata within these adjacent catchments seem to have diverged relatively recently, roughly 130 000 years ago. Contemporary dispersal seems to be absent between catchments but we suggest that climate fluctuations during the Pleistocene resulted in extensive floods that promoted historical movement of aquatic organisms across catchment boundaries.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 1–16.     

Spatiotemporal analysis of gene flow in Chesapeake Bay Diamondback Terrapins (Malaclemys terrapin)

There is widespread concern regarding the impacts of anthropogenic activities on connectivity among populations of plants and animals, and understanding how contemporary and historical processes shape metapopulation dynamics is crucial for setting appropriate conservation targets. We used genetic data to identify population clusters and quantify gene flow over historical and contemporary time frames in the Diamondback Terrapin (Malaclemys terrapin). This species has a long and complicated history with humans, including commercial overharvesting and subsequent translocation events during the early twentieth century. Today, terrapins face threats from habitat loss and mortality in fisheries bycatch. To evaluate population structure and gene flow among Diamondback Terrapin populations in the Chesapeake Bay region, we sampled 617 individuals from 15 localities and screened individuals at 12 polymorphic microsatellite loci. Our goals were to demarcate metapopulation structure, quantify genetic diversity, estimate effective population sizes, and document temporal changes in gene flow. We found that terrapins in the Chesapeake Bay region harbour high levels of genetic diversity and form four populations. Effective population sizes were variable. Among most population comparisons, estimates of historical and contemporary terrapin gene flow were generally low (m ≈ 0.01). However, we detected a substantial increase in contemporary gene flow into Chesapeake Bay from populations outside the bay, as well as between two populations within Chesapeake Bay, possibly as a consequence of translocations during the early twentieth century. Our study shows that inferences across multiple time scales are needed to evaluate population connectivity, especially as recent changes may identify threats to population persistence.