Mitochondrial DNA analysis of the genetic relationships among populations of scad mackerel (Decapterus macarellus, D. macrosoma, and D. russelli) in South-East Asia

The genetic relationships among South-East Asian populations of the scad mackerels Decapterus macarellus, D. macrosoma and D. russelli (Pisces: Carangidae) were investigated. In 1995 and 1996, 216 fish were sampled in seven localities spanning the seas of Indonesia and were examined for restriction-site polymorphisms using ten restriction enzymes for the mitochondrial (mt) DNA control region, amplified by the polymerase chain-reaction. The inferred phylogeny of haplotypes led to the recognition of three distinct mitochondrial lineages or phylads consistent with the distinctions of current taxonomy. All 15 mtDNA haplotypes found in D. macarellus and all 9 haplotypes found in D. macrosoma were arranged as star-like clusters, suggesting recent evolutionary history. In contrast, the phylad formed by 6 haplotypes in D. russelli from the Sulawesi Sea exhibited diffuse topology, suggesting that ancestral lineages of this species have been retained to the present. Average nucleotide-divergence estimates between haplotypes of different phylads were between 0.042 and 0.135, suggesting ancient separation, in consistency with published allozyme data. High levels of haplotype diversity, but no geographical heterogeneity, was detected within D. macarellus from the Molucca Sea and the Banda Sea. Populations of D. macrosoma exhibited both significant differences between adjacent regions (Sunda Strait and Java Sea), and broadscale genetic homogeneity from the South China Sea to the Sulawesi Sea via the Java Sea and Makassar Strait. The geographic isolation of the D. macrosoma population sampled in the Sunda Strait suggests that this region constitute a sharp transition zone between the Indian Ocean and the Sunda Shelf. Near-monomorphism of haplotypes and low nucleotide diversity (d _X) were observed in the samples of D. macrosoma from the continental shelf (haplotype-diversity estimates, h, = 0.00 to 0.25 ± 0.08 and d _X = 0.000 to 0.002). This was in contrast to the comparatively high haplotype and nucleotide diversities observed in other pelagic fish species including D. macarellus (h = 0.82 ± 0.05, d _X = 0.012 to 0.015) and D. russelli (h = 0.63 ± 0.12, d _X = 0.016), and in the oceanic D. macrosoma population sampled in the Sunda Strait (h = 0.67 ± 0.31, d _X = 0.005). We hypothesise that this may be the consequence of recent and perhaps repeated bottleneck events that have affected the D. macrosoma population sampled on the continental shelf. Received: 29 September 1997 / Accepted: 3 September 1999     

The distribution and habitat preference of coastally occurring minke whales (Balaenoptera acutorostrata) in the outer southern Moray Firth, northeast Scotland

The coastal waters of the Moray Firth in northeast Scotland (57o41’N 2o40’W) provide rich, inshore feeding grounds for minke whales (Balaenoptera acutorostrata) during the summer and autumnal months. In order to better understand the habitat selection, movements and feeding ecology of the animals utilising this North Sea region, distribution data from the southern coastline of the outer Moray Firth were subsequently examined with respect to the marine physiography of the area, specifically the environmental variables water depth, slope, aspect and sediment-type. A total of 305 minke whale encounters – collected from dedicated boat surveys conducted between May and October 2001 to 2006 inclusive – were used in the construction of a Geographic Information System (GIS) for the 860 square-km study site. The subsequent analysis revealed a strong spatial preference by whales in this location for water depths between 20 and 50 metres (mean 46.9 m, SD = 30.9), steep slopes (mean 75.7 degrees, SD = 8.9), a northerly-facing aspect and sandy-gravel sediment type. Kruskal-Wallis tests for variance confirmed that the distribution of B. acutorostrata was significantly different across each of these physiographic features examined (P < 0.05). In particular, water depth and sediment type were shown to be highly correlated with the frequency of whales observed (Spearman’s Rank Correlation P < 0.05 for depth and sediment respectively). From these results, we conclude that sea bottom characteristics may be used to predict the fine-scale distribution of minke whales on their feeding grounds; the physiographic features identified providing valuable proxies for inferring prey distributions in the absence of fisheries data. However, an appreciation of both abiotic and biotic factors (using a combination of GIS and remote sensing outputs) is clearly desirable for ecosystem-based management approaches for the coastal conservation of these whales. The application of GIS capacities to ecological studies based largely on field data of these marine mammals is highly recommended in the present study to cetologists, environmental modellers and conservation managers alike.     

Home range, activity and distribution patterns of a temperate rocky-reef fish, Cheilodactylus fuscus

Red morwong (Cheilodactylus fuscus; Cheilodactylidae: Perciformes), are large temperate rocky reef fish that are patchily distributed in local aggregations of 3 to >100 fish. The home ranges and aggregating behaviour of red morwong were investigated using external tags (n = 114, over 35 mo) and ultrasonic transmitters (n = 9 over 20 d) at coastal and estuarine sites in south-east Australia. Adult red morwong had a mean home range during the day of 1865 m² (SE = 268). Night movement, determined by ultrasonic telemetry, indicated a significantly greater mean home range of 3639 m² (SE = 416). Home range did not differ significantly with size or sex. Fish movements were greatest and aggregation sizes varied most during the non-reproductive mid-summer period. Tagged fish (N = 20) displaced 200 to 900 m returned in 1 to 3 d to the point of capture, often traversing open sand habitat and other aggregations. During the day, fish were commonly found on bouldered habitat. During the night, these site-specific aggregations fragmented as fish dispersed over a variety of substrata, with crepuscular peaks in activity. Therefore, estimates of habitat-specificity and feeding patterns collected only by day may give misleading results. Received: 28 June 1996 / Accepted: 8 July 1998     

Remote Antarctic feeding ground important for east Australian humpback whales

Understanding the dynamics of population recovery is particularly complex when an organism has multiple, remote breeding and feeding grounds separated by one of the longest known migration routes. This study reports on the most comprehensive assessment of humpback whale (Megaptera novaeangliae) movements between remote Antarctic waters south of New Zealand and east Australia (EA), and the migratory corridors and breeding grounds of Australia and Oceania. A total of 112 individual whales were identified; 57 from microsatellites and 61 by fluke with 23 % (n = 26) matched to sites outside Antarctica. Despite large datasets from other southern regions being included in the comparison, the whales were predominantly linked to EA (n = 24). Only two matches to the Oceania catalogues directly north was surprising; therefore the primary feeding grounds of these endangered whales still remain unknown. The confirmation of the Balleny Islands as an important feeding ground for EA whales could provide an insight into reasons behind the rapid recovery of this population. Determining the feeding grounds of Oceania’s whales may explain whether prey energetics or migration length are limiting factors to their recovery and will allow an understanding of future ecosystem changes in these whales.     

Post-glacial population history and genetic structure of the northern clingfish (Gobbiesox maeandricus), revealed from mtDNA analysis

Mitochondrial d-loop sequences were analyzed to characterize the phylogeographic and population genetic structure of the northern clingfish (Gobbiesox maeandricus Girard). Sequence analysis of 378 bp from 111 individuals sampled in 14 localities along the northeast Pacific coast and within the Strait of Georgia from 1996 to 1999 revealed marked genetic differentiation (Φ_ct=0.247) among regional population groupings. The gene genealogy distinguished two major clades of haplotypes separated by at least 1.1% sequence divergence. One clade with very low haplotype diversity (h=0.2095, n=18) occurred only within the recently unglaciated Strait of Georgia. The other clade had high haplotype diversity (h=0.8808, n=93) and was found in all populations. High haplotype diversity was found in open coastal populations, both north and south of the maximum extant of the Wisconsin ice sheet, suggesting that the clingfish range was not pushed to a southern refugium during the last glacial maximum. A nested clade analysis also did not detect a large northward expansion from a single southern refugium. The level of sequence divergence and coalescent-based analyses suggest that the observed patterns of polymorphism are the result of Pleistocene diversification within multiple refugia, followed by population expansion and asymmetrical lineage introgression. Received: 5 February 2000 / Accepted: 31 August 2000     

Temporal stability and spatial divergence of mitochondrial DNA haplotype frequencies in red drum (Sciaenops ocellatus) from coastal regions of the western Atlantic Ocean and Gulf of Mexico

Restriction-site variation in mitochondrial (mt) DNA was assayed among 1675 red drum (Sciaenops ocellatus Linnaeus) sampled from 20 localities along the southeastern coast of the USA (western Atlantic) and the Gulf of Mexico (Gulf). Up to four consecutive year-classes (cohorts) were sampled at most localities. Nucleotide-sequence divergence among 170 mtDNA haplotypes identified ranged (in percentage) from 0.184 to 1.913, with a mean (±SD) of 0.887 ± 0.300. Comparisons of mtDNA haplotype frequencies across year-classes within localities were non-significant, indicating temporal stability of breeding components within localities. Significant heterogeneity in mtDNA haplotype frequencies was found across all localities, between (pooled) samples from the western Atlantic and the Gulf, and among geographically spaced, regional groupings in the Gulf. Genetic divergence between subpopulations of red drum in the western Atlantic and Gulf follows a pattern exhibited in other marine fishes, and probably stems from physical (historical environmental heterogeneity, absence of suitable habitat, and current patterns) and, perhaps, behavioral factors. Genetic differences among red drum in the Gulf appear to be due largely to an isolation-by-distance effect that is attributable to behavioral factors. The latter may include female philopatry to natal bays or estuaries, limited offshore (coastwise) movement of females relative to their natal bay or estuary, or both. Genetic divergence among red drum in the Gulf occurs despite high gene flow (estimated as the number of genetic effective migrants in an island mode). Conservation and management of red drum should be based on the premise that strategies for a given bay or estuary will impact geographically proximal bays or estuaries more than distal ones. Trajectories of correlograms in spatial autocorrelation analysis suggest a geographic neighborhood size, relative to genetic migration of red drum from a bay or estuary, of roughly 500 to 600 km. Received: 22 July 1998 / Accepted: 19 November 1998     

Phylogeography of the ocean quahog (Arctica islandica): influences of paleoclimate on genetic diversity and species range

The ocean quahog, Arctica islandica (Linnaeus, 1767), is a commercially important bivalve found on continental shelves throughout much of the North Atlantic. To assess genetic subdivision in this species, we sequenced 385 nucleotides of the mitochondrial cytochrome b (cyt b) gene from 83 specimens collected from 12 localities between September 1998 and July 1999 (based on preliminary data, the Internal Transcribed Spacers, ITS, of the nuclear ribosomal repeat were not useful). The cyt b data delimited 11 haplotypes with 0.26 to 8.1% nucleotide difference (coded by 36 variable nucleotide positions) among them. Only three haplotypes were detected in 39 specimens collected along the USA coastline, compared to five haplotypes from nine Icelandic individuals. The western Atlantic populations ranging from Penobscot Bay (Maine, USA) to southern Virginia showed relatively low diversity and appeared genetically similar in that region. Based on the presence of shared haplotypes, AMOVA analyses, and phylogenetic reconstructions, Icelandic populations appear to be more genetically similar to western Atlantic populations than eastern Atlantic populations. Specimens from the Faroe Islands (n=4) show mixed affinities. These data are consistent with the hypothesis that a warm Holocene climatic optimum (ca. 7,500 years BP), and not glacial refugia, shaped the present-day genetic structure in A. islandica. Received: 18 January 2000 / Accepted: 26 June 2000     

Genetic population structure of southern bluefin tuna (Thunnus maccoyii )

Samples of southern bluefin tuna, Thunnus maccoyii (Castelnau), taken from off the coasts of South Africa, Western Australia, South Australia and Tasmania from 1992 to 1994 were analysed for six polymorphic allozyme loci (ADA ^*, GDA ^*, GPI-A ^*, MPI ^*, PGDH ^* and PGM-1 ^*, n = 595 to 733 per locus) and for mitochondrial DNA variants revealed by three restriction enzymes (Bam HI, Bcl I and Eco RI) detecting polymorphic cut sites (n = 555). No significant spatial heterogeneity was detected. There were no sex-related differences in allele or mtDNA haplotype frequencies. Juveniles (30 to 35 cm and 46 to 54 cm) from what are thought to be two temporally-separated spawning peaks showed no significant genetic differentiation. There were also no significant differences in allele or haplotype frequencies between fish smaller than 70 cm and those larger than 70 cm. These data are consistent with the null hypothesis of a single unit stock of southern bluefin tuna, with a single spawning area. This is located to the south of Java and off the north-west coast of Australia. Received: 28 August 1996 / Accepted: 30 September 1996     

Effects of organochlorines on the ultrastructure of the liver of the damselfish Parma microlepis from reefs in New South Wales, Australia

  Parma microlepis (Günther) were collected from Malabar, an urban location close to the centre of Sydney, Australia, and from Jervis Bay, a reference location 170 km south of the city centre. At each location, fish were collected from two sites separated by 100 to 200 m. The ultrastructure of normal liver tissue is described based on 20 female fish collected from Jervis Bay, where fish are known to be exposed to low levels of organochlorine contaminants. Alterations in the endoplasmic reticulum, mitochondria, lysosomes and nuclei of hepatocytes were identified and quantified in the liver tissue of fish from this location and compared to alterations in 20 female fish collected from Malabar, where fish are exposed to higher concentrations of organochlorine pesticides such as DDT compounds. There were significant differences in the percentage of hepatocytes with swollen mitochondria (F = 124.025, df = 2, 2, P = 0.008) and atypical nuclei (F = 22.198, df = 2, 2, P = 0.043) between sites (100 to 200 m apart), but there were no clear differences between the percentage of structural alterations in the hepatocytes of P. microlepis from Jervis Bay and Malabar. Associations between liver morphology and the organochlorines aldrin, dieldrin, DDE and chlordane were examined using a Pearson correlation matrix. Significant correlations were detected between the percentage of hepatocytes with dilated endoplasmic reticulum and the concentrations of the pesticide aldrin (r = 0.600, df = 11, r _{crit(α = 0.05)} = 0.553). Significant associations were also detected between the percentage of hepatocytes with disorganised endoplasmic reticulum and the concentrations of dieldrin and DDE residues in fish (r = 0.576, r = 0.567, respectively, df = 13, r _{crit (α = 0.05)} = 0.514). However, there was little evidence that ultrastructural alterations in fish responded to increasing concentrations of these pesticides in a consistent dose-response manner. Received: 20 October 1998 / Accepted: 24 November 1999     

Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata

Nucleotide variation in cytochrome c oxidase subunit I (COI) was used to examine population structure in three invasive bryozoans: Bugula neritina (Linnaeus, 1758), Watersipora subtorquata (d’Orbigny, 1852), and W. arcuata (Banta, 1969). These species are found on ship hulls and have a short (≤2 days) larval phase. Samples were collected from 1998–2001 at multiple sites in Australia, and in Hong Kong, New Zealand, Hawaii, California, Curaçao, and England. B. neritina is known to include three cryptic species, including species Type S (Davidson and Haygood in Biol Bull 196:273–280, 1999) which occurs on the east and west coasts of the USA. One haplotype recorded previously in the USA, S1, was found to be widespread, occurring throughout Australia and in Hong Kong, Curaçao, Hawaii, and England. W. subtorquata, a Caribbean–Atlantic species which has invaded southern Australia, New Zealand, and California, had low nucleotide diversity in these areas (π=0.0016±0.0014), consisting of three haplotypes connected by one or two nucleotide mutations. W. arcuata, an Eastern-Pacific native, had comparatively high diversity (π=0.0221±0.0115) in introduced populations from Australia and Hawaii. In each species, identical haplotypes were identified on separate coastlines providing evidence of widespread, rather than genetically independent, introductions. The major contrast in nucleotide diversity suggests that different propagule-source models explain introductions.     

Variability in abundances of fishes associated with seagrass habitats in relation to diets of predatory fishes

The spatial, diel and tidal variability in the abundance of piscivorous fishes and their teleost prey, and the dietary composition of predatory fishes were investigated in beds of Heterozostera tasmanica within Port Phillip Bay, Australia, from September 1997 to February 1998. Predatory and prey fish assemblages were sampled from beds of H. tasmanica at three locations during each combination of diel (day and night) and tidal (high and low) cycles. Pelagic and benthic crustaceans represented >60% by abundance of the diets of all predatory fishes. Seven species, 54% of all predatory fishes, were piscivorous. These piscivores consumed individuals from seven families, 36.8% of the fish families being associated with seagrass. Western Australian salmon, Arripis truttacea (Arripidae) (n = 174) and yank flathead, Platycephalus speculator (Platycephalidae) (n = 46) were the most abundant piscivores. A. truttacea consumed larval/post-larval atherinids, gobiids and sillaginids. P. speculator consumed late-juvenile/adult atherinids, clinids and gobiids. While the abundances of piscivores varied between locations (P < 0.001) and diel periods (P = 0.028), the relative differences in piscivore abundance between sites and diel periods were not consistent between tides. The abundances of A. truttacea varied in a complex way amongst sites, diel period and tidal cycle, as shown by a three-way interaction between these factors (P = 0.026). Only during diurnal periods at St. Leonards was the abundance of A. truttacea significantly higher during high than low tides (P < 0.001). During the other diel periods at each site, the abundance of A. truttacea did not vary. P. speculator was significantly more abundant nocturnally (P = 0.017). The abundance of small (prey) fishes varied significantly amongst sites (P < 0.001). During the day, the abundance of small fishes did not vary between high and low tides (P = 0.185), but their nocturnal abundance was greater during low tide (P < 0.001). Atherinids (n = 1732) and sillaginids (n = 1623) were the most abundant families of small fishes. Atherinids were significantly more abundant nocturnally (P = 0.005) and during low tides (P = 0.029), and varied significantly amongst sites (P < 0.001). Sillaginids varied significantly only amongst sites (P < 0.001). Seagrass beds provide a foraging habitat for a diverse assemblage of predatory fishes, many of which are piscivorous. Anti-predator behaviour and amongst-location variability in abundances of piscivorous fishes may explain some of the diel and tidal, and broad-scale spatial patterns in small-fish abundances. Received: 23 July 1999 / Accepted: 18 January 2000     

Genetic structure,lack of sex-biased dispersal and behavioral flexibility in the pair-living fat-tailed dwarf lemur, <Emphasis Type="Italic">Cheirogaleus medius</Emphasis>

Mating system and dispersal patterns influence the spatio-genetic structure within and between populations. Among mammals, monogamy is rare, and its socio-genetic consequences have not been studied in detail before. The goal of our study was to investigate population history, demographic structure, and dispersal patterns in a population of pair-living fat-tailed dwarf lemurs, Cheirogaleus medius, a small, nocturnal primate from western Madagascar, and to infer their underlying behavioral mechanisms. Tissue samples for DNA extraction were obtained from a total of 140 individuals that were captured in two subpopulations about 3 km apart. Analyses of mtDNA variability at the population level revealed very low levels of genetic variability combined with high haplotype diversity, which is indicative of a recent population bottleneck. We found no evidence for spatial clustering of same-sexed individuals with identical haplotypes within each of two subpopulations but significant clustering between them. Thus, a high level of local subpopulation differentiation was observed (F _ST = 0.230). The sexes showed equal variances in the number of individuals representing each haplotype, as well as equal levels of aggregation of identical haplotypes. Hence, both sexes disperse from their natal area, one pattern expected in a pair-living mammal. There is a possibility of behavioral and social flexibility in this species, however, because we documented pronounced differences in density and sex ratio between the two subpopulations, suggesting that single study sites or populations may not be representative of a given local population or even species.     

Need and nestmates affect begging in tree swallows

We conducted an experiment on nestling tree swallows (Tachycineta bicolor) to examine predictions from signalling models for the evolution of conspicuous begging behaviour. Specifically, we examined the relationship between (1) nestling begging intensity and hunger, (2) begging intensity and parental provisioning and (3) begging intensity and nestmate condition. Forty broods of 9-day-old nestlings were removed from their nests for 1 h and assigned to one of the following three treatments: (1) all nestlings in the brood deprived of food (n = 13), (2) all nestlings in the brood fed (n = 11) or (3) half the nestlings in the brood deprived and half fed (n = 16). Videotapes before and after the treatments showed that begging intensity increased in broods in which all of the nestlings had been deprived and decreased in broods in which all of the nestlings had been fed. Deprived nestlings in the half-and-half treatment did not change their begging intensity in response to treatment, while fed nestlings in this treatment group showed a decrease in begging intensity. Parent tree swallows increased their feeding rate to deprived broods and decreased their rate to fed broods. Within broods, parents decreased their feeding rate to fed nestlings, but showed no significant change in feeding to deprived nestlings. Our results suggest that begging intensity is influenced by hunger and that parents appear to respond to variation in begging intensity. The begging of nestmates also appears to influence begging independently of need. These results are consistent with predictions derived from signalling models of begging. Received: 20 June 1997 / Accepted after revision: 19 January 1998     

Mitochondrial DNA variation in king mackerel (Scomberomorus cavalla) from the western Atlantic Ocean and Gulf of Mexico

King mackerel (Scomberomorus cavalla Cuvier) collected in 1992 and 1993 from 13 localities along the Atlantic coast of the southeastern USA and in the northern Gulf of Mexico were surveyed for variation in mitochondrial (mt)DNA and a nuclear-encoded dipeptidase locus (PEPA-2). Both polymorphic and fixed mtDNA restriction sites were identified and mapped using conventional and polymerase chain-reaction (PCR)-based methods. Heterogeneity in mtDNA haplotype frequencies was found only in comparisons of pooled haplotypes from Atlantic localities versus pooled haplotypes from Gulf localities. This finding indicates weak genetic divergence between king mackerel from the Atlantic and those from the Gulf. Frequencies of two PEPA-2 alleles essentially paralleled previous findings: one allele (PEPA-2a) was common among samples from western Gulf localities, whereas the other allele (PEPA-2b) was common among samples from Atlantic and eastern Gulf localities. There was considerable variation in PEPA-2 allele frequencies within broadly-defined regions. Variation in mtDNA haplotypes and PEPA-2 genotypes was independent, as was variation in mtDNA haplotypes with sex or age of individuals. Variation in PEPA-2 genotypes was not independent of sex or age of individuals. The latter result suggests that frequencies of PEPA-2 alleles in samples of king mackerel may stem, in part, from sex and age distributions of individuals within samples, and indicates that caution should be exercised in using allelic variation at PEPA-2 as a measure of population (stock) structure in king mackerel. The discordance in spatial patterning of mtDNA haplotypes versus PEPA-2 alleles across the Gulf (i.e. homogeneity in mtDNA haplotype frequencies versus heterogeneity in PEPA-2 allele frequencies) may be due to either female excess at several localities, sex-biased migration, or both. Observed patterns of genetic variation also are consistent with the hypothesis that king mackerel in the western Atlantic may have been subdivided during Pleistocene glaciation, and that the current distribution of PEPA-2 alleles may be a historical artefact. Received: 17 December 1996 / Accepted: 2 April 1997     

Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Population genetic and phylogeographic structure of wahoo,<Emphasis Type="Italic"> Acanthocybium solandri</Emphasis>, from the western central Atlantic and central Pacific Oceans

The wahoo, Acanthocybium solandri (Cuvier, 1832), is a pelagic, highly migratory, scombroid fish, distributed worldwide throughout tropical and warm temperate seas. To evaluate population genetic and phylogeographic structure against a null hypothesis of panmixia, the entire mitochondrial DNA control-region (~890 base pairs) was sequenced for 231 wahoo. Samples were collected from 1997 to 2001 from seven sites: North Carolina (NC; n=23), east central Florida (CF; n=30), Bimini, Bahamas (BB; n=40), southern tip of Florida (SF; n=21), Cayman Islands (CI; n=23), northern Gulf of Mexico (NG; n=54), and Hawaii (HI; n=40). Inter-annual samples were obtained from four of these locations (NC, BB, SF, NG). Seventeen haplotypes were shared by individuals within and among samples; 187 singleton haplotypes were observed. Within-sample haplotype diversities ranged from 0.995 to 1.000 (overall h=0.999) and within-sample nucleotide diversities ranged from 0.049 to 0.055 (overall =0.053). A neighbor-joining tree based on inter-haplotypic distances revealed two monophyletic lineages differing by 13.6% nucleotide divergence. Nested within each major lineage were several, well-supported subclades. There was no evidence of temporal heterogeneity in haplotype distributions. Partitioning mtDNA variation, 99.75% of the variance was within samples and 0.25% (P=0.307) between samples; the fixation index (_ST=0.0025) was not significant. Likewise, pairwise _ST values were low or negative, and none were significant on a table-wide basis. Exact tests for sample differentiation in haplotypes were also non-significant. All population analyses were consistent with the null hypothesis of panmixia. However, analytical power was limited by sample size. Mismatch distributions were inconsistent with expected distributions based on sudden-expansion and static-growth models. Wahoo exhibit concurrently high haplotype and nucleotide diversities, presumably a consequence of secondary contact between historical subpopulations rather than a long, stable evolutionary history. Given the level of geographic and individual sampling, wahoo thus far represent the sole example of a scombroid or xiphioid fish exhibiting coarse-grain genetic homogeneity across a broad, inter-oceanic range despite a deeply coalescing genealogical structure. Accordingly, cooperative fishery management on a broad, inter-ocean scale may be warranted.Communicated by J.P. Grassle, New Brunswick     

Sociality,Bateman’s gradients,and the polygynandrous genetic mating system of round-tailed ground squirrels (<Emphasis Type="Italic">Xerospermophilus tereticaudus</Emphasis>)

Historically, most mammals have been classified as polygynous; although recent molecular evidence suggests that many mammals may be polygynandrous, particularly the ground-dwelling sciurids. We genotyped 351 round-tailed ground squirrels (Xerospermophilus tereticaudus) using seven microsatellite loci to determine paternity in 31 litters from 2004 to 2007. Polygyny was evident in all years except in 2007, when the population size was reduced. Multiple paternity occurred in the majority of litters (55%) with 2.5 ± 0.26 sires/litter (n = 31). Forty-nine percent of resident males (n = 114) sired offspring, and of males that sired offspring (n = 56) 27% sired young in multiple litters in a single breeding season. Litter size was positively correlated with the number of sires. Through an indirect analysis of paternity, we found 21 litters (68%) with an average relatedness of 0.5 or less. Males had a greater opportunity for sexual selection (I _s = 1.60) than females (I _s = 0.40); Bateman’s gradient was also greater in males (1.07 ± 0.04, n = 56) than females (0.82 ± 0.08, n = 31). The mating system in round-tailed ground squirrels defined through genetic analyses and Bateman’s gradients is polygynandrous compared to the previously suggested polygynous mating system as established by behavioral observations and fits within the predictions of the ground squirrel sociality models. Upon evaluating the predictions of the sociality models among sciurid species, we found a negative relationship between the level of sociality with litter size and the average percentage of multiple paternity within a litter. Thus, recent genetic information and reclassification of mating systems support the predictions of the ground-dwelling squirrel sociality models.     

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups. ‘Resident’-type (fish-eating) killer whales were more abundant than the ‘transient’-type (mammal-eating). Abundance estimates of resident killer whales (991 [95% CI = 379–2,585] [IGS] and 1,587 [95% CI = 608–4,140] [PEGS]), were at least four times greater than those of the transient killer whales (200 [95% CI = 81–488] [IGS] and 251 [95% CI = 97–644] whales [PEGS]). The IGS estimate of abundance is preferred for resident killer whales because the estimate based on PEGS data may show an upward bias. The PEGS estimate of abundance is likely more accurate for transients. Residents were most abundant near Kodiak Island in the northern Gulf of Alaska, around Umnak and Unalaska Islands in the eastern Aleutians, and in Seguam Pass in the central Aleutians. This ecotype was not observed between 156 and 164°W, south of the Alaska Peninsula. In contrast, transient killer whale sightings were found at higher densities south of the Alaska Peninsula between the Shumagin Islands and the eastern Aleutians. Only two sightings of ‘offshore’-type killer whales were recorded during the surveys, one northeast of Unalaska Island and the other south of Kodiak Island. These are the first estimates of abundance of killer whale ecotypes in the Aleutian Islands and Alaska Peninsula area and provide a baseline for quantifying the role of these top predators in their ecosystem. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.

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Karyotypic characterization of the great sturgeon, Huso huso, by multiple staining techniques and fluorescent in situ hybridization

A karyotype analysis by several staining techniques was carried out on the great sturgeon, Huso huso (Linnaeus, 1758). The karyotype (2n = 118 ± 2) was composed of 42 pairs of meta-/submetacentric chromosomes and 17 pairs of acrocentrics/microchromosomes. Constitutive heterochromatin was mainly located at the centromeric regions of the acrocentric chromosomes. The biarmed chromosomes showed weak C-bands. Fluorescent staining with GC-specific chromomycin A₃ showed clearly recognizable fluorescent regions, whereas a more uniform base composition was revealed by the AT-specific 4,6-diamidino-2 phenylindole. After Ag-staining, nucleolar organizer regions could be observed on the short arms of two medium-sized submetacentrics and on two acrocentrics. Digoxigenated 28S and 5S rDNA probes, prepared from Acipenser naccarii DNA and hybridized to metaphase chromosomes, showed signals on six and two chromosomes, respectively. The telomeric sequence (TTAGGG) _n detected by FISH was located at both ends of each chromosome. Results are discussed in relation to karyotype organization and evolution in sturgeons. Received: 1 April 1998 / Accepted: 21 July 1998     

Oxygen transport by the hemocyanin of the protobranch mollusc Solemya reidi

Oxygen transport by the hemocyanin of the protobranch mollusc Solemya reidi Bernard was studied in native hemolymph samples. Clams were collected from two different reducing environments, beneath log booms in Alberni Inlet, British Columbia, Canada, and from the sewage effulent in Santa Monica Bay, California, USA. The hemocyanin concentration in a pooled hemolymph sample (n = 10 individuals) was 33.5 mg ml⁻¹. The mean hemolymph pH of five Alberni clams maintained for 3 wk in mud was 7.96 ± 0.06. No significant variation in hemocyanin oxygen-affinity or cooperativity was found for hemocyanin in whole hemolymph samples from these five individually studied clams. There was a significant difference only at 15 °C in the oxygen affinity of hemocyanin in pooled whole-hemolymph samples from S. reidi collected from Alberni Inlet compared with clams collected from Santa Monica Bay. Little effect of temperature on hemocyanin oxygen-affinity was found for temperatures below 20 °C; above 20 °C, the oxygen affinity was reduced for clams from both sites. Temperature and pH had no apparent affect on hemocyanin cooperativity. Moderate Bohr shifts were found at all temperatures examined. The presence of physiologically relevant concentrations of thiosulphate in hemocyanin samples resulted in a decrease in hemocyanin oxygen-affinity, opposite to the effect on hemocyanin found for the hydrothermal vent crab Byth-ograea thermydron, but thiosulphate had no effect on hemocyanin cooperativity. Received: 2 September 1997 / Accepted: 6 February 1998