Molecular insight into the population structure of common and spotted dolphins inhabiting the pelagic waters of the Northeast Atlantic

Several cetacean species exhibit fine-scale population structure despite their high dispersal capacities and the apparent continuity of the marine environment. In dolphins, most studies have focused on coastal areas and continental margins, and they revealed differentiated populations within relatively small geographic areas, sometimes in conjunction with a specialisation for different habitats (ecotypes). We analysed the population genetic structure of short-beaked common dolphins (Delphinus delphis) and Atlantic spotted dolphins (Stenella frontalis) in the Azores and Madeira, the two most isolated archipelagos of the North Atlantic. The archipelago of the Azores is divided into three groups of islands and stands 900 km away from Madeira. It is not known whether individuals migrate between groups of islands and archipelagos, nor whether distinct ecotypes are present. These questions were investigated by genetic analyses of 343 biopsy samples collected on free-ranging dolphins. The analyses consisted in sequencing part of the mitochondrial hyper-variable region, screening up to 14 microsatellite loci, and molecular sexing. Results did not unravel any population structure at the scale of the study area. Lack of differentiation matches expectations for spotted dolphins, which are transient in both archipelagos, but not for common dolphins, which are present year-round in the Azores and potentially resident. Absence of genetic structure over hundreds and even thousands of kilometres implies the existence of gene flow over much larger distances than usually documented in small delphinids, which could be achieved through individual movements. This finding indicates that population structure in oceanic habitat differs from that observed in coastal habitat.     

New insights on population genetic structure of <Emphasis Type="Italic">Delphinus delphis</Emphasis> from the northeast Atlantic and phylogenetic relationships within the genus inferred from two mitochondrial markers

The taxonomic status of common dolphins (Delphinus sp.) remains controversial despite the increased number of studies focusing on its populations. Two species are presently recognized, Delphinus delphis and D. capensis. Apart from a phylogeographic study of the genus Delphinus, genetic studies focusing specifically in the northeast (NE) Atlantic remain scarce. Following ecological and morphological evidence for the existence of different common dolphin morphotypes in the Portuguese coast, we examined the population structure of D. delphis from the NE Atlantic by comparing DNA sequences from two mitochondrial regions (control region and cytochrome b gene). Additionally, we compared the sequences obtained with existing sequences of D. delphis from the Azores, Black Sea, Canary Islands, Pacific Ocean, D. capensis and also two closely related delphinid species (Stenella coeruleoalba and Tursiops truncatus). In the analysis of the NE Atlantic populations, we found evidence for the existence of some level of genetic differentiation. In the broader phylogenetic analysis, D. delphis and D. capensis did not show reciprocal monophyly and we found a group of highly divergent individuals. We discuss the possibility for the existence of two divergent lineages that have evolved independently, a separate subspecies or events of introgressive hybridization. These findings could have important implications on a taxonomic level, although further investigation based on a larger geographical scale and on nuclear loci information will certainly elucidate the origin of these highly divergent individuals.     

Population differentiation and taxonomic status of the exploited limpet Patella candei in the Macaronesian islands (Azores,Madeira, Canaries)

There has been considerable confusion in the taxonomy of limpets of the North East Atlantic Ocean and Mediterranean Sea, particularly those from the Macaronesian islands. The present study compared populations of the intertidal limpet Patella candei d'Orbigny from the Azores, Madeira and the Canaries with those of P. caerulea Linnaeus and P. depressa Pennant from the European and African continental coasts. No major differences in radular morphology were detected between the three species. However, electrophoretic analysis of 15 enzyme loci gave overall genetic identity (I) values of 0.5 between the three species, indicating that they cannot be regarded as conspecific as previously thought, and suggesting that P. candei is endemic to the Macaronesian islands. Comparisons of P. candei within these islands showed that, although populations did not differ with respect to radular morphology and soft-body parts, populations from the Azores were distinct from those in Madeira and the Canaries in shell shape and gene frequencies. Individuals from the Azores had, no average, taller shells and longer radulae, while those in Madeira and the Canaries had a shallow, depressed and stellate shell form. This was interpreted as being due to the wider habitat distribution of the species in the Azores compared to Madeira and the Canaries. Electrophoretic results showed that P. candei from the Azores differed from P. candei in Madeira and the Canaries by almost 40% of the loci investigated (I=0.660), suggesting that the former is a separate endemic species. An I value of 0.969 between populations in Madeira and the Canaries was typical of conspecific populations.     

Tropical fishes in a temperate sea: evolution of the wrasse <Emphasis Type="Italic">Thalassoma pavo</Emphasis> and the parrotfish <Emphasis Type="Italic">Sparisoma cretense</Emphasis> in the Mediterranean and the adjacent Macaronesian and Cape Verde Archipelagos

The northeastern Atlantic and the Mediterranean Sea share geological histories and display great faunal affinities. The majority of the Mediterranean species have Atlantic origins, with a few species with tropical affinities. These include the parrotfish Sparisoma cretense and the wrasse Thalassoma pavo that are restricted to the subtropical northeastern Atlantic, the Macaronesian archipelagos (Azores, Madeira, and Canaries) and the southern Mediterranean. The Pleistocene glaciations have been described as having different effects on the fauna of the two regions. During glacial peaks, Mediterranean waters remained warmer than those of the adjacent Atlantic. Within the eastern Atlantic, the effects of Pleistocene glaciations were differentiated. Here, we perform a comparative analysis focusing on T. pavo and S. cretense populations from the northeastern Atlantic and the Mediterranean to assess the effects of Pleistocene glaciations in these two species. Sequences from the mitochondrial control region were obtained and analyzed combining phylogeographic and demographic approaches. Gene flow between Atlantic and Mediterranean populations was shown to be very high. The Mediterranean populations of T. pavo and S. cretense showed high levels of genetic diversity, even in the eastern basin, pointing to an ancient colonization event. This suggests that both species must have been able to persist in the Mediterranean during the cold Pleistocene periods. Historical migration estimates revealed a Mediterranean towards Atlantic trend in the case of T. pavo, which may reflect the re-colonization of areas in the Atlantic by fish that survived the cold phases in relatively warmer Mediterranean refugia. Our data also showed that within the Macaronesian Archipelagos, migrations occurred from Madeira towards the Azores, for both T. pavo and S. cretense, thus supporting a post-glacial colonization of the Azores by fish that persisted in the warmer region of Madeira. Similar geographic distributions, thermal affinities, and means of dispersion for T. pavo and S. cretense resulted in a similar response to the effects of Pleistocene glaciations, as evidenced by identical phylogeographic patterns.     

Completely disjunct mitochondrial DNA haplotype distribution without a phylogeographic break in a planktonic developing gastropod

Planktonic developing organisms are generally assumed to be good dispersers showing little genetic structuring in neutral markers. At first glance, this also applies to the planktonic developing periwinkle Tectarius striatus, an endemic gastropod from Macaronesia (i.e. Azores, Madeira, Canary Islands and Cape Verde Islands), where the only sign of genetic structuring hitherto is provided by a non-significant allozyme/RAPD heterogeneity between the Cape Verde Islands and the other archipelagos. However, partial sequences of the mitochondrial cytochrome b and cytochrome oxidase I genes now show that the Cape Verde Islands and the three other archipelagos have no haplotypes in common, whereas the latter three do share several haplotypes. Nevertheless, this highly disjunct haplotype distribution does not entail a phylogeographic break separating the haplotypes of both areas in two reciprocally monophyletic groups. This remarkable geographic and phylogenetic structuring may be explained by assuming that T. striatus colonized the Macaronesian archipelagos in periods when sea levels were lower (and/or volcanic activity was higher), so that seamounts peaked above sea level and could act as stepping-stones. Yet, after the last glacial period seamounts submerged, thus preventing further stepping-stones mediated dispersal of T. striatus between the Cape Verde Islands and the other archipelagos, while not affecting dispersal among the latter because of their closer proximity and connectivity. Hence, these contrasting patterns of neutral genetic variation in T. striatus show that genetic structuring in planktonic developing species may be far more complex than is usually assumed.     

Population differentiation of the Atlantic spotted dolphin (Stenella frontalis) in the western North Atlantic, including the Gulf of Mexico

Information about the genetic population structure of the Atlantic spotted dolphin [Stenella frontalis (G. Cuvier 1829)] in the western North Atlantic would greatly improve conservation and management of this species in USA waters. To this end, mitochondrial control region sequences and five nuclear microsatellite loci were used to test for genetic differentiation of Atlantic spotted dolphins in the western North Atlantic, including the Gulf of Mexico (n=199). Skin tissue samples were collected from 1994–2000. Significant heterozygote deficiencies in three microsatellite loci within samples collected off the eastern USA coast prompted investigation of a possible Wahlund effect, resulting in evidence for previously unsuspected population subdivision in this region. In subsequent analyses including three putative populations, two in the western North Atlantic (n=38, n=85) and one in the Gulf of Mexico (n=76), significant genetic differentiation was detected for both nuclear DNA (R _ST=0.096, P≤0.0001) and mitochondrial DNA (Φ _ST=0.215, P≤0.0001), as well as for all pair-wise population comparisons for both markers. This genetic evidence for population differentiation coupled to known biogeographic transition zones at Cape Hatteras, North Carolina and Cape Canaveral, Florida, USA, evidence of female philopatry, and preliminary support for significant genetic differences between previously documented morphotypes of Atlantic spotted dolphins in coastal and offshore waters all indicate that the biology and life history of this species is more complex than previously assumed. Assumptions of large, panmictic populations might not be accurate in other areas where S. frontalis is continuously distributed (e.g., eastern Atlantic), and could have a detrimental effect on long-term viability and maintenance of genetic diversity in this species in regions where incidental human-induced mortality occurs.

---

Distribution of common dolphins (Delphinus spp.) in the western Atlantic Ocean: a critical re-examination

Due to indications that misidentification (largely confusion among dolphins of the genera Delphinus and Stenella) in the past had led to erroneous assumptions of distribution of the two species of common dolphins (Delphinus delphis and D. capensis) in the western Atlantic Ocean, we conducted a critical re-examination of records of the genus Delphinus from this region. We compiled 460 ‘plottable’ records, required support for confirmation of genus and species identifications, and found many records lacking (and some clearly misidentified). When we plotted only the valid records (n = 364), we found evidence of populations in only three areas, and apparent absence throughout much of the tropical/subtropical regions. Off the east coast of the US and Canada, D. delphis is found from the Georgia/South Carolina border (32°N) north to about 47–50°N off Newfoundland. Since the 1960s, they have apparently been absent from Florida waters. There is no evidence that dolphins of the genus occur in the Gulf of Mexico. Reports of common dolphins from most of the Caribbean Basin are also rejected, and the only place in that region where they are confirmed to occur is off central-eastern Venezuela (a coastal D. capensis population). Off eastern South America, common dolphins appear to be restricted to south of 20°S. There is a coastal long-beaked population found in the South Brazil Bight, and one or more short-beaked populations south and offshore of this (ranging south to at least northern Argentina). The results are very different from commonly-accepted patterns of distribution for the genus in the Atlantic. Most areas of distribution coincide with moderate to strong upwelling and common dolphins appear to avoid warm, tropical waters. This study shows that great care must be taken in identification of similar-appearing long-beaked delphinids, and that uncritical acceptance of records at face value can lead to incorrect assumptions about the ranges of the species involved.     

Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Population structure and connectivity of the European conger eel (Conger conger) across the north-eastern Atlantic and western Mediterranean: integrating molecular and otolith elemental approaches

Genetic variation (mtDNA) of the European conger eel, Conger conger, was compared across five locations in the north-eastern Atlantic (Madeira, Azores, South Portugal, North Portugal and Ireland) and one location in the western Mediterranean (Mallorca). Genetic diversity of conger eel was high, and differentiation among regions was not significant. Additionally, comparisons of element:Ca ratios (Sr:Ca, Ba:Ca, Mn:Ca and Mg:Ca) in otolith cores (larval phase) and edges (3?months prior to capture) among the Azores, North Portugal, Madeira and Mallorca regions for 2?years indicated that variation among regions were greater for edges than cores. Therefore, while benthic conger may display residency at regional scales, recruitment may not necessarily be derived from local spawning and larval retention. Furthermore, data from otoliths suggest a separated replenishment source for western Mediterranean and NE Atlantic stocks. The combination of genetics and otolith chemistry suggests?a population model for conger eel involving a broad-scale dispersal of larvae, with limited connectivity for benthic juvenile life stages at large spatial scales, although the existence of one or multiple spawning grounds for the species remains uncertain.     

Fine-scale population structure of estuarine bottlenose dolphins (Tursiops truncatus) assessed using stable isotope ratios and fatty acid signature analyses

Stable isotope ratios and fatty acid signature analyses were employed to examine the fine-scale population structure of a year-round resident population of 600–800 bottlenose dolphins (Tursiops truncatus) in the Indian River Lagoon (IRL), Florida. The IRL, a 250-km-long estuary running along the central east coast of Florida (28.0°N, 80.6°W), is comprised of the northern and southern IRL, Mosquito Lagoon (ML), Banana River (BR), and St. Lucie Estuary. Samples of skin and blubber were collected from dead stranded (n = 61, 1994–2004) and live dolphins (n = 153, 2002–2007, 2010, 2011) from throughout the IRL and surrounding environs. Using stable isotopes (SI), dolphins could be assigned to a ML subpopulation, a St. Lucie Estuary subpopulation, and an IRL subpopulation. Fatty acid signature analysis (FASA) allowed for finer resolution, detecting ML and BR subpopulations, a separation of northern and southern IRL subpopulations, and a St. Lucie Estuary subpopulation. Differences between sexes were detectable within subpopulations using FASA, but not using SI. This may indicate that males and females are foraging in similar locations at a similar trophic level (detected using SI), but are varying in the types or proportions of specific prey (indicated by FASA). The combination of these complementary analyses results in a powerful tool for assessing fine-scale population substructure.     

Winter diet shift of long-beaked common dolphins (Delphinus capensis) feeding in the sardine run off KwaZulu-Natal, South Africa

Unpredictable inter-annual variations in the timing, spatial extent and intensity of the sardine run (Sardinops sagax) have been documented in recent years along the coastline of KwaZulu-Natal, South Africa. This study aimed to determine whether variations in the availability of sardine were reflected in the diet and condition of common dolphins (Delphinus capensis) over the past three decades. Stomach contents from 95 common dolphins incidentally caught between 2000 and 2009 were analysed and compared to historical data collected between 1972 and 1992. A shift in the principal prey species consumed by the dolphins was observed over the past 30 years. Prior to 1992, sardine comprised up to 49 % of the total stomach contents by mass, whilst chub mackerel (Scomber japonicus) was the dominant prey recorded in the stomach contents between 2000 and 2009 (66 %). As common dolphins prey on locally abundant fish species, this dietary shift may indicate changes in the availability of the most abundant fish prey. Stable isotope analyses of tooth tissue revealed no significant changes in the δ¹⁵N and δ¹³C isotopic ratios over the past three decades (P = 0.283 for N and P = 0.922 for C). Blubber thickness as a percentage of body length and blubber weight as a percentage of total body weight were assessed as indicators of animal condition. No significant changes in proportional blubber weight or blubber thickness were seen over the last 40 years (1970–2009) for all age cohorts. This species appears to be well adapted to cope with changes in prey species availability, without impacting on body condition.     

Fine-scale genetic structure in short-beaked common dolphins (<Emphasis Type="Italic">Delphinus</Emphasis><Emphasis Type="Italic">delphis</Emphasis>) along the East Australian Current

Oceanographic processes play a significant role in shaping the genetic structure of marine populations, but it is less clear whether they affect genetic differentiation of highly mobile vertebrates. We used microsatellite markers and mtDNA control region sequences to investigate the spatial genetic structure of short-beaked common dolphins (Delphinus delphis) in southeastern Australia, a region characterised by complex oceanographic conditions associated with the East Australian Current (EAC). A total of 115 biopsy samples of dolphins were collected from six localities spanning approximately 1,000 km of the New South Wales (NSW) coastline. We found evidence for contrasting genetic diversity and fine-scale genetic structure, characterised by three genetically differentiated populations with varying levels of admixture. Spatial genetic structure was not explained by a model of isolation by distance, instead it coincides with main patterns of oceanographic variation along the EAC. We propose that common dolphins along the EAC may be adapted to three water masses recently characterised in this region.     

Population structure of short-beaked common dolphins (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the North Atlantic Ocean as revealed by mitochondrial and nuclear genetic markers

The understanding of population structure and gene flow of marine pelagic species is paramount to monitoring, management and conservation studies. Such studies are often hampered by the potentially high dispersal behavior of the species, the lack of obvious geographical barriers in the marine environment and the scarce sample availability. Short-beaked common dolphins (Delphinus delphis) are widespread in coastal and open-ocean habitats of the North Atlantic Ocean, nevertheless population structure and migratory patterns are poorly understood. Furthermore, concern has been raised about the status of the species because large numbers of dolphins have been taken incidentally in several fisheries throughout the North Atlantic in the past decades. In the present study, a large number of individual samples were obtained from seasonal and spatial aggregations of common dolphins from western (wNA) and eastern North Atlantic (eNA) regions, mostly using opportunistic sampling (i.e. from incidental entanglement in fishing gear or beach-cast carcasses). Genetic variability was investigated using nuclear (14 microsatellite loci) and mitochondrial (360 bp of the control region) genetic markers. Levels of genetic diversity were relatively high in all sampled areas and no evidence of recent reduction of effective population size (i.e. bottleneck) was detected at the nuclear loci. Significant population structure was detected between the two main regions (wNA and eNA) where it appeared to be more pronounced at mitochondrial (F _ST = 0.018, P < 0.001) than nuclear markers (F _ST = 0.005, P < 0.05), indicating the presence of at least two genetically distinct populations of common dolphins in the North Atlantic Ocean. In contrast, no significant genetic structure was detected between temporal aggregations of dolphins from within the same region, suggesting possible seasonal movement patterns at a regional scale. The observed levels of genetic differentiation between classes of markers are discussed here as a possible consequence of migratory patterns or recent population subdivision. An erratum to this article can be found at     

Major histocompatibility complex class II variation in bottlenose dolphin from Adriatic Sea: inferences about the extent of balancing selection

The bottlenose dolphin (Tursiops truncatus) is the most common cetacean species worldwide and the only marine mammal species resident in the Croatian part of the Adriatic Sea. To gain insight into genetic diversity of bottlenose dolphins at adaptively important loci relevant to conservation, we analysed the polymorphism of major histocompatibility complex (MHC) genes, which play a key role in pathogen confrontation and clearance. Specifically, we examined the diversity of MHC class II DRA, DQA and DQB alleles in 50 bottlenose dolphins from the Adriatic Sea collected between 1997 and 2011 and in 12 animals from other Mediterranean locations. Notable variation in DQA, DQB and three-locus haplotypes was found, with all 10 DQA and 12 DQB alleles encoding unique protein products. Analysis of the ratio of non-synonymous to synonymous substitution rates suggests that positive selection acts at both highly variable loci. Phylogenetic analyses revealed trans-species polymorphism at the DQB locus, strongly indicating the influence of balancing selection in the long term. In fact, the balancing selection observed in bottlenose dolphins is higher than that reported for most other cetaceans and comparable to that seen in terrestrial mammals.     

Pregnancy patterns of pantropical spotted dolphins (Stenella attenuata) in the eastern tropical Pacific determined from hormonal analysis of blubber biopsies and correlations with the purse-seine tuna fishery

The northeastern offshore population of the pantropical spotted dolphin (Stenella attenuata) in the eastern tropical Pacific remains listed as depleted under the Marine Mammal Protection Act, and recent estimates of abundance show that recovery has been slow. One hypothesis for the slow recovery is that continued chase and encirclement by the tuna fishery negatively affects reproduction. Insufficient life-history sampling in this region over the last two decades makes traditional estimates of reproductive rates impossible. Here, we examine the current reproductive patterns of these dolphins by measuring blubber progesterone (BP) concentrations in biopsy samples to assess pregnancy state. BP was quantified in 212 biopsies from female offshore spotted dolphins sampled between 1998 and 2003 in the northeastern tropical Pacific, and we found that 11.5 % of the biopsied females (mature and immature) were pregnant. The relationship between pregnancy and fishery exposure was analyzed, and we found that pregnant females were exposed to significantly less fishery activity than non-pregnant ones (p = 0.022), suggesting that the fishery may have an inhibitive effect on pregnancy. Spatial analysis indicated that pregnancy was more aggregated than random (p < 0.05) at a scale up to 180-nmi, with the highest proportion pregnant in the mouth of the Gulf of California, an area with relatively low reported fishery activity.     

Population structure of red porgy, <Emphasis Type="Italic">Pagrus pagrus</Emphasis>, in the Atlantic Ocean

The red porgy, Pagrus pagrus (L.), is a protogynous sparid associated with reefs and hard bottom habitat throughout the warm-temperate Atlantic Ocean. In this study, the degree of geographic population differentiation in Atlantic populations was examined with microsatellite and mitochondrial DNA markers (mtDNA). Six microsatellite loci were amplified and scored in 690 individuals from the eastern North Atlantic (Crete, Madeira, and Azores), western North Atlantic (North Carolina to Florida, and the eastern Gulf of Mexico), and Brazil. At two loci, fixed allelic differences were found among the three major geographic areas, while frequency differences were observed at three other loci. The DNA of 371 individuals was amplified at the mtDNA control region, and 526 bp were sequenced. Tamura–Nei’s D was used as a measure of nucleotide diversity and divergence: diversity averaged 0.011 within samples, while the corrected divergence averaged 0 between samples within the same area and 0.061 between samples from different areas. Transversion haplotype minimum spanning networks, nucleotide divergence, and F _ST values all show that the western Atlantic samples were more closely related to each other than any was to samples from the eastern North Atlantic. Within the western North Atlantic, no significant population differentiation was observed, and within the eastern North Atlantic, only the Azores sample showed detectable differences from Crete and Madeira. These data indicate general homogeneity within large areas, and deep divisions between these areas. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Reproduction in short-beaked common dolphins (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) from the western North Atlantic

A single population of short-beaked common dolphins (Delphinus delphis Linnaeus) occurs in the western North Atlantic from Florida to Newfoundland. Dolphins killed in a swordfish driftnet fishery between 1989 and 1998 provided samples for the present study. These were combined with additional observations from carcasses collected from stranding programs. Samples were examined from 74 female and 161 male short-beaked common dolphins. A highly significant male bias in the sex ratio of both the bycatch and stranding samples suggests that sex-based habitat partitioning or school segregation occurs in this population. Age was estimated from decalcified thin sections of teeth and testis and ovarian samples were examined both macro- and microscopically. Analyses showed that reproduction was both seasonal and synchronized. Females reached sexual maturation at approximately age eight; males matured at 9.5 years. Conception occurred during July and August (mean day 24 July). Gestation was estimated to last just under a year, so most females gave birth during July and August (mean day 15 July). Annual pregnancy rate was estimated to be between 25 and 33%. Males undergo a fivefold seasonal increase in testes mass and a twofold increase in seminiferous tubule diameter. Peak sperm production potential was observed in July. At the end of the breeding season testis size and activity decreased after August. Male dolphins in this population are hypothesized to engage in some form of sperm competition.     

Prey selection by resident common bottlenose dolphins (tursiops truncatus) in Sarasota Bay, Florida

Prey selection was investigated in wild, resident common bottlenose dolphins, Tursiops truncatus, during the summer months in Sarasota Bay, Florida, USA. Stomach content analyses of 15 dolphins with extensive sighting histories and well-documented distributions were used to determine prey use. Prey availability was assessed by purse seine surveys. We compared the relative abundances of prey available to estimates of prey use at closely matching spatial and temporal scales. G-tests determined that dolphins in this study significantly selected for prey at the species, family, and soniferous/non-soniferous prey levels (G _adj  = 753.98–1,775.93, df = 1–21, p ≤ 0.01). While comprising only 6.3% of the total available prey, soniferous fishes accounted for 51.9% of the total prey consumed. Manly’s standardized forage ratios and 95% Bonferroni confidence intervals determined significant positive selection for soniferous prey and against non-soniferous prey (β^S = 0.9461 vs. β^NS = 0.0539). Dolphins selected against Gerridae, Clupeidae, and Sparidae (β ≤ 0.0014), as well as against all the species within those families (β ≤ 0.0190). It is likely that passive listening for soniferous prey provides an ecological or energetic advantage to cetaceans utilizing this specific foraging technique.     

Genetic variation in the neritic squid Loligo forbesi (Myopsida: Loliginidae) in the northeast Atlantic Ocean

Horizontal starch gel electrophoresis was employed to investigate levels of genetic differentiation between 13 samples of the neritic squid species Loligo forbesi Steenstrup obtained from throughout the majority of its known geographical range. Six enzyme loci identified in a preliminary study as being polymorphic were screened for variation between samples. No significant differences in allele distribution were detected between any of the samples obtained from the Faroe Bank in the north to Lisbon in the south, suggesting that squid throughout this range in the vicinity of the continental shelf are able to maintain panmixia, and effectively belong to a single population sharing a common gene pool. No clinal variation in allele distribution was detected throughout this range, a result which complements the findings of a detailed morphological companion study of the same individuals. Comparison of this homogenous European continental shelf population with squid from the Azores revealed highly significant (P<0.01) differences in allele distribution at five of the six polymorphic enzyme loci studied. A genetic identity value (I) equivalent to 0.93 over 33 loci was obtained. Analysis of F-statistics suggested migration rates between sites to be as low as one individual per five generations, a rate deemed insufficient under most models to prevent divergence by random genetic drift. The large distance and oceanic depths separating the Azores from continental Europe seem to present an effective barrier to gene flow to L. forbesi, a squid belonging to a family considered to be confined in distribution to relatively shallow, near coastal waters. The two populations of squid in the Azores and along the European continental shelf currently both ascribed to L. forbesi should therefore probably best be regarded as relative subspecies.