Molecular and morphometric variation in European populations of the articulate brachiopod Terebratulina retusa

Molecular and morphometric variation within and between population samples of the articulate brachiopod Terebratulina spp., collected in 1985–1987 from a Norwegian fjord, sea-lochs and coastal sites in western Scotland, the southern English Channel (Brittany) and the western Mediterranean, were measured by the analysis of variation in the lengths of mitochondrial DNA (mtDNA) fragments produced by digestion with nine restriction endonucleases and by multivariate statistical analysis of six selected morphometric parameters. Nucleotide difference within each population sample was high. Nucleotide difference between population samples from the Scottish sites, both those that are tidally contiguous and those that appear to be geographically isolated, were not significantly different from zero. Nucleotide differences between the population samples from Norway, Brittany, Scotland and the western Mediterranean were also very low. Morphometric analysis confirmed the absence of substantial differentiation.     

Genetic structure of summer flounder (Paralichthys dentatus) populations north and south of Cape Hatteras

Genetic divergence among samples of summer flounder (Paralichthys dentatus, Linnaeus) was estimated to evaluate the effect of Cape Hatteras, a well-known zoogeographic barrier in the western Atlantic Ocean, on intraspecific gene flow. Previous studies based on meristic and morphometric variation and tagging data suggested that gene flow might be limited among summer flounder populations north and south of Cape Hatteras, although the high vagility of both juveniles and adults suggests otherwise. We analyzed genetic diversity revealed in the mitochondrial DNA (mtDNA) control region in samples of juveniles and adults collected in 1992 to 1996 from coastal sites from Buzzard's Bay, Massachusetts to Charleston, South Carolina. In contrast to previous morphological studies, analyses of mtDNA variation reveal no significant population subdivision centered around Cape Hatteras. Received: 17 September 1997 / Accepted: 8 September 1998     

The Mytilus edulis species complex in southwest England: effects of hybridization and introgression upon interlocus associations and morphometric variation

Loglinear analysis of electrophoretic data from two hybrid Mytilus edulis x galloprovincialis populations in southwest England revealed non-significant associations between genotypes at four allozyme loci, each of which is partially diagnostic for differences between the two taxa. Significant non-random genotypic associations within the context of the non-significant model involved all four assayed loci equally, consistent with their occurrence in a relatively tight linkage group. Multivariate analyses were used to examine electrophoretic variation from the two hybrid populations, and morphometric variation in the hybrid populations and in four allopatric (two M. edulis and two M. galloprovincialis) populations from western Europe While the number of hybrid mussels is high at both sites (22% at Croyde, 53% at Whitsand) the two taxa have largely maintained the genetic differences which exist between them in allopatry. However, morphological differences between the taxa have been eroded for mussels within the hybrid zone, whereas these differences are quite pronounced for mussels from allopatric populations. It is proposed that each taxon within the genus maintains its genetic identity, despite high dispersal potential, widespread hybridization, and high levels of introgression, as a result of adaptation to different environments. The worlwide occurrence of all four Mytilus hybrid zones at ecotones between recognized biogeographical provinces which are characterized by differences in temperature and salinity is consistent with such an interpretation.     

Allozyme heterozygosity and escape response performance of the scallops, <Emphasis Type="Italic">Argopecten purpuratus</Emphasis> and <Emphasis Type="Italic">Placopecten magellanicus</Emphasis>

Multilocus allozyme heterozygosity (MLH) has been positively correlated with growth in some marine bivalves and was suggested to facilitate swimming activity in pectinids. Using two highly mobile scallops, Placopecten magellanicus and Argopecten purpuratus, we examined escape response performance and morphometric characteristics as a function of allelic variability at metabolic loci. Ten allozyme systems were used for A. purpuratus and 7 for P. magellanicus. In each species, the morphometric characteristics and escape response parameters were analyzed separately using principal components analysis (PCA) and the scores of the major principal components were related to allozyme heterozygosity. In both P. magellanicus and A. purpuratus, positive correlations were found between MLH and morphometric parameters, but escape response parameters were only positively linked to MLH in P. magellanicus, and then weakly. The hypothesis that MLH improves fitness of pectinids by increasing the capacity to escape predators is not supported.     

Intraspecific Chromosome Number Variation: a Neglected Threat to the Conservation of Rare Plants

Abstract: The effectiveness of rare plant conservation will increase when life history, demographic, and genetic data are considered simultaneously. Inbreeding depression is a widely recognized genetic concern in rare plant conservation, and the mixing of genetically diverse populations in restoration efforts is a common remedy. Nevertheless, if populations with unrecognized intraspecific chromosome variation are crossed, progeny fitness losses will range from partial to complete sterility, and reintroductions and population augmentation of rare plants may fail. To assess the current state of cytological knowledge of threatened and endangered plants in the continental United States, we searched available resources for chromosome counts. We also reviewed recovery plans to discern whether recovery criteria potentially place listed species at risk by requiring reintroductions or population augmentation in the absence of cytological information. Over half the plants lacked a chromosome count, and when a taxon did have a count it generally originated from a sampling intensity too limited to detect intraspecific chromosome variation. Despite limited past cytological sampling, we found 11 plants with documented intraspecific cytological variation, while 8 others were ambiguous for intraspecific chromosome variation. Nevertheless, only one recovery plan addressed the chromosome differences. Inadequate within‐species cytological characterization, incomplete sampling among listed taxa, and the prevalence of interspecific and intraspecific chromosome variation in listed genera, suggests that other rare plants are likely to have intraspecific chromosome variation. Nearly 90% of all recovery plans called for reintroductions or population augmentation as part of recovery criteria despite the dearth of cytological knowledge. We recommend screening rare plants for intraspecific chromosome variation before reintroductions or population augmentation projects are undertaken to safeguard against inadvertent mixtures of incompatible cytotypes.     

Evidence that reef-wide patterns of coral bleaching may be the result of the distribution of bleaching-susceptible clones

The hypothesis that intraspecific variation in coral bleaching is a result of the distribution of bleaching-susceptible clonal genotypes (genets) was addressed using photoquadrats recorded during the 1987 Caribbean bleaching event on a reef dominated by Montastraea annularis (Morphotype I), together with manipulative experiments with Porites porites. Nearest-neighbor analysis showed that bleached colonies (ramets) of M. annularis at 10 m depth had a high probability (0.80) of having a nearest bleached neighbor of the same genet rather than a bleached ramet of a different genet. Furthermore, the frequency distributions of bleached ramets of M. annularis in the photoquadrats was significantly different from a Poisson distribution, suggesting that bleached ramets were aggregated on the reef. Manipulative experiments with P. porites from 15 m depth showed that some genets were more susceptible to thermal bleaching than others, since three genets had significantly different rates of zooxanthellae loss when exposed to elevated temperatures in tanks receiving irradiances similar to those found in situ. These results suggest that the in situ patchy distribution of bleached ramets could correspond to the distribution of certain genets, and that adjacent genets can exhibit sufficiently different phenotypes to account for intraspecific variation in bleaching. Further studies of genet-specific coral bleaching may provide valuable insights into the causes and consequences of bleaching.     

Morphological and DNA sequence variation in the kelp Costaria costata (Phaeophyta)

Kelp species show substantial intraspecific morphological variation. The annual kelp, Costaria costata (C. Agardh) Saunders, is found in the lower intertidal and subtidal regions of shores which vary in exposure from strongly wave-exposed to sheltered. Phenotypic variation was studied in this monotypic genus. Individuals of C. costata were collected from two morphologically-distinct populations (one wave-exposed and the other wave-sheltered) at Cape Beale and Stanley Park, Vancouver, Canada, in May 1985. Analysis of morphological variation, using multivariate statistical techniques, indicated significant differentiation between plants from the two sites. Blade widths and thicknesses were the major discriminating variables. Restriction fragment length polymorphism (RFLP) analysis of nuclear DNA with 42 anonymous probes demonstrated one polymorphism. Most of the probes are hypothesized to encode highly repeated, dispersed sequences. These repeat sequence probes comprised 80.9% of all cloned fragments. Primary sequence analysis of 1595 base pairs of small-subunit ribosomal DNA and 204 base pairs of upstream sequence (probe pCc 18) failed to show any divergence between plants from the two sites.     

Geographic variation in Loligo forbesi in the Northeast Atlantic Ocean: analysis of morphometric data and tests of causal hypotheses

Geographic variation in the squid Loligo forbesi was investigated using multivariate analysis of morphometric and meristic characters in samples of squid taken from 13 localities in the Northeast Atlantic Ocean. Two character sets, body morphometrics and beak morphometrics, indicated similar patterns of variation, with squid from the Azores differing markedly from those on the continental shelf. No consistent pattern was apparent in meristic data. Partial Mantel tests indicated that similarity matrices for morphological data were significantly correlated with distance matrices for (a) geographic proximity, (b) whether the capture site was on the continental shelf or the Azorean bank, and (c) (beak data only) average seasurface temperature at site of capture. Partial Mantel tests on allozyme data for the same individuals support hypothesis (b). The results suggest that L. forbesi in the Azores may reasonably be regarded as a distinct stock, differing significantly from L. forbesi on the continental shelf.     

Biodiversity Consequences of Genetic Variation in Bark Characteristics within a Foundation Tree Species

Abstract: The developing field of community genetics has the potential to broaden the contribution of genetics to conservation biology by demonstrating that genetic variation within foundation plant species can act to structure associated communities of microorganisms, invertebrates, and vertebrates. We assessed the biodiversity consequences of natural patterns of intraspecific genetic variation within the widely distributed Australian forest tree, Eucalyptus globulus. We assessed genetic variation among geographic races of E. globulus (i.e., provenances, seed zones) in the characteristics of tree‐trunk bark in a 17‐year‐old common garden and the associated response of a dependent macroarthropod community. In total, 180 macroarthropod taxa were identified following a collection from 100 trees of five races. We found substantial genetically based variation within E. globulus in the quantity and type of decorticating bark. In the community of organisms associated with this bark, significant variation existed among trees of different races in composition, and there was a two‐fold difference in species richness (7–14 species) and abundance (22–55 individuals) among races. This community variation was tightly linked with genetically based variation in bark, with 60% of variation in community composition driven by bark characteristics. No detectable correlation was found, however, with neutral molecular markers. These community‐level effects of tree genetics are expected to extend to higher trophic levels because of the extensive use of tree trunks as foraging zones by birds and marsupials. Our results demonstrate the potential biodiversity benefits that may be gained through conservation of intraspecific genetic variation within broadly distributed foundation species. The opportunities for enhancing biodiversity values of forestry and restoration plantings are also highlighted because such planted forests are increasingly becoming the dominant forest type in many areas of the world.     

Morphologically plastic responses to shading in the zoanthids Zoanthus sansibaricus and Palythoa tuberculosa

Zoanthids of the order Zoantharia (Anthozoa: Hexacorallia) exhibit high intraspecific morphological variability, but whether this is due to polymorphism, phenotypic plasticity or a combination of both remains unknown. To address this knowledge gap, in November 2010, eight colonies each of Zoanthus sansibaricus and Palythoa tuberculosa were sampled from three reefs off the south of mainland Singapore and transplanted to a shallow experimental site. The colonies were then distributed under two types of treatment frames: control and shaded. After 87 days, morphometric characters were extracted from macro-images. Reaction norms, principal components analysis, analysis of variance and canonical discriminant analysis all demonstrated light-induced changes in morphology. Patterns of plastic changes were similar for both species: shaded colonies had larger polyps as compared to control colonies. The presence of plastic responses in zoanthids may facilitate their colonization of a broad range of habitats as well as help them to withstand temporal changes in their environment.     

Identification of Illex coindetii, I. illecebrosus and I. argentinus (Cephalopoda: Ommastrephidae) throughout the Atlantic Ocean; by body and beak characters

Shortfinned squid species of the genus Illex support commercial fisheries throughout the Atlantic Ocean and Mediterranean Sea. Previous identification of interspecific and intraspecific populations by morphological and size-at-maturity studies have not provided conclusive results. We analysed morphometric body and beak variables (24 characters) in three species of the genus (I. coindetii, I. illecebrosus and I. argentinus), using a geographic and seasonal series of 33 populations for 1,500 specimens of I. coindetii, I. illecebrosus and I. argentinus. Residuals of the regression between each morphometric body and beak variable and mantle length were used as input in a stepwise discriminant analysis. Species discrimination by body and hectocotylus characters required at least eight variables and resulted in high correct-classification percentages for I. coindetii and I. argentinus (75% and 90%, respectively), whereas the best identification resulted from beak characters (83% correctly classified). Size of the suckerless basal arm, sucker-bearing length and beak lateral wall discriminated best among I. coindetii from northern Iberia, northwest Iberia (year-1996) and Ireland in the Atlantic and western Mediterranean versus middle and eastern Mediterranean samples. Canadian shelf and American samples were discriminated from Canadian slope I. illecebrosus. Winter/shelf and winter/slope samples of I. argentinus seemed to form a single biological group separated from Falkland Island, 46°S/autumn spawners and 46°S/1996 specimens along the Patagonian Shelf. No significant sexual or maturity polymorphism was obtained. Discriminant analysis optimised population diagnosis on a morphometric basis of interest in fisheries strategies. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0796-7.     

Population genetic analysis of Ensis directus unveils high genetic variation in the introduced range and reveals a new species from the NW Atlantic

We report current genetic variation of populations of the razor shell Ensis directus (Conrad 1843) (Mollusca: Bivalvia: Pharidae) in native (North American) and introduced (European) ranges using nuclear and mitochondrial sequence-based markers. We expected less variation within the introduced range, especially considering the frequent mass mortality events observed in Europe since the species was recorded for the first time in 1978. However, we found higher variation in Europe. The possible significance of temporal fluctuations of genetic variation, limited effect of random genetic drift, and multiple introductions are discussed. Interestingly, the multiple-introduction hypothesis contrasts with the gradual colonisation of European coastal waters but is supported by trained clustering analysis and by the intensity of transatlantic shipping. Genetic and morphometric evidence strongly supports that examined individuals from a supposed E. directus population from Newfoundland (Canada) belong to a separate species. This new Ensis is formally described here and named E. terranovensis n.sp.     

Visual associative learning in two desert ant species

Visual cues are important navigational tools for many solitary foraging insects. In addition to information provided by path integration, desert ants learn and use visual cues for homing back to their nest. In this study, we compared the visually based learning of two desert ant species: the North African Cataglyphis fortis and the Australian Melophorus bagoti, each of which lives in ecologically similar but visually different environments. In our experiment, ants’ choice performance was measured by training foragers in a channel system. We used a decision box with two visual stimuli during their homebound trips, with one of the stimuli always being the correct one that provided thoroughfare. To determine any habitat effects on learning, we examined intraspecific comparisons in C. fortis with different nest surroundings. The intraspecific comparison in C. fortis revealed no differences in learning the task. In general, C. fortis showed little learning in our task and the results were similar for ants from barren and cluttered environments. Overall, M. bagoti learned the task faster and had a higher level of accuracy than C. fortis. One explanation for this species-specific difference could be that the cluttered habitat of M. bagoti favours the evolution of visual associative learning more so than the plain habitat of C. fortis.     

Diet and respiration of the small planktonic marine copepod Oithona nana

The functional responses of Oithona nana (Giesbr.) to various phytoplankton and zooplankton food species are described. The food species were divided into three size categories, the seasonal abundances of which were measured in Loch Turnaig, a Scottish sea loch in 1977. The seasonal variations in feeding rates in the sea for each size class were derived. The seasonal variation in respiration rate of O. nana was measured, and metabolic requirements were claculated as between 6 and 40% of the food material estimated as being eaten. O. nana differs from other common copepods in having a wide food-particle size spectrum and a low metabolic rate. It is suggested that these adaptations constitute the strategy whereby O. nana maintains its population levels throughout the year.     

Lack of population structure in the fiddler crab Uca annulipes along an East African latitudinal gradient: genetic and morphometric evidence

Phylogeographic patterns in Uca annulipes sampled from 30 locations across an East African latitudinal gradient were investigated using mitochondrial DNA cytochrome oxidase I sequences and analysed together with patterns of morphometric differentiation. Four hundred and four specimens along the east African coast were sampled, and 18 haplotypes were encountered. Haplotype and nucleotide diversity values were very low and the phylogenetic analysis did not reveal any clear phylogeographic structure. Furthermore, the analysis of molecular variance and pairwise Φ _ST values showed no significant spatial population differentiation. Mismatch analyses and tests of neutrality supported the hypothesis that this species has undergone a fairly recent demographic expansion. Our results, therefore, failed to demonstrate significant geographical structure in the pattern of genetic variation, indicating that populations of U. annulipes are capable of extensive gene flow among mangroves along the coast. The genetic structure of this species could be panmictic due to a high amount of gene flow along the geographical gradient in study during the planktonic larval phase, when larvae are carried along stream by the dominant currents. Moreover, the morphometric analysis performed did not reveal differences of shape differentiation according to a geographical pattern, although significant differences among the sampling areas were found.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Allozymic and morphometric evidence for two stocks of the European anchovy Engraulis encrasicolus in Adriatic waters

Allozyme electrophoresis and morphometric analyses were used to investigate the stock structure of European anchovies (Engraulis encrasicolus L.) captured by Italian vessels in the Adriatic Sea. Twenty four putative enzyme loci were studied, all of which exhibited genotypic proportions in accordance with Hardy-Weinberg predictions. Two loci, IDHP-2 ^* (isocitrate dehydrogenase, 1.1.1.42) and G3PDH-2 ^* (glycerol-3-phosphate dehydrogenase, 1.1.1.8), showed significant allele-frequency differences among samples. IDHP-2 ^* 100 frequencies ranged between 0.35 and 0.88, with lower frequencies recorded in northern waters and areas close to the Italian coastline, while G3PDH-2 ^* 100 frequencies followed a similar, if less distinct, pattern. For both loci, the trends observed were spatially stable over a 2 yr sampling period. A putative stock-boundary was superimposed onto a map of IDHP-2 ^* 100 frequencies, and a bootstrapped dendrogram confirmed the genetic separation of the two putative stocks, one located around the north-western Adriatic and the other in the central-southern region. Canonical variate analysis (CVA) of morphometric data collected using a truss network indicated that the two putative stocks were morphologically distinct; 89.6 and 88.3% of north-western and central-southern anchovies, respectively, were correctly assigned by discriminant-function analysis. This variation may be related to the presence of two anchovy colour phenotypes, silver and blue, in the Adriatic Sea. Silver anchovies are characteristic of northern areas, while the larger blue fish are found mainly in the deeper southern waters. Current assessment models for the fishery are based on the concept of a consistent identity between consecutive catches at single ports, which our data reject. We discuss the possibility of partitioning fishing effort based on the proportions of the two stocks landed at individual ports.     

Queen acceptance and the complexity of nestmate discrimination in the Argentine ant

In most social insect species, individuals recognize and behave aggressively towards non-nestmate conspecifics to maintain colony integrity. However, introduced populations of the invasive Argentine ant, Linepithema humile, exhibit pronounced variation in intraspecific aggression denoting diversity in nestmate recognition behavior, which possibly shapes their social structure and the varying levels of unicoloniality observed among these populations. One approach to better understand differential aggression behaviors towards conspecifics and recognition cue perception and response in L. humile is to examine variation in nestmate discrimination capability among genetically distinct colonies under different social contexts. Consequently, we investigated the dynamics of queen and worker recognition in southeastern US L. humile queenless and queenright colonies by measuring rates of non-nestmate worker and queen adoption and intercolony genetic similarity. Aggression levels between colony pairs differed and were associated with non-nestmate worker, but not queen adoption. Adoption of queens and workers was a function of host colony origin, while colony queen number affected adoption of queens, but not workers, with queens more readily accepted by queenless hosts. Fecundity of adopted non-nestmate queens was comparable to that of rejected non-nestmate and host colony queens, suggesting that queen fecundity did not affect adoption decisions. Genetic similarity between colonies ranged from 30 to 77% alleles shared, with more genetically similar colonies showing lower levels of intraspecific aggression. Non-nestmate queens and workers that were more genetically similar to host colony workers were more likely to be adopted. We provide the first evidence for the role of L. humile colony queen number on queen discrimination and suggest an effect of resident queens on worker conspecific acceptance thresholds. Our findings indicate a role for genetically based cues in L. humile nestmate recognition. However, subtle discrimination capability seems to be influenced by the social context, as demonstrated by more frequent recognition errors in queenless colonies.     

Lipid composition and partitioning of deepwater chondrichthyans: inferences of feeding ecology and distribution

The composition of lipids and fatty acids was determined for the livers, muscle, pancreas, kidney and stomach fluids of deepwater chondrichthyan species (including 11 squaliformes, 3 chimaeriformes, 1 hexanchiforme and 3 carcharhiniformes) caught as bycatch from continental waters off south-eastern Australia. The lipid class, fatty acid and fatty alcohol composition differed markedly in each tissue and in each species. The lipid and fatty acid composition of large, lipid-rich (38–70% wet weight, ww) livers demonstrated the multifunctional role of this organ in: lipid distribution, storage and biosynthesis, and buoyancy regulation. In the liver, the importance of certain lipids (including squalene, diacylglyceryl ethers, triacylglycerols and to a lesser extent wax esters) as mediators of buoyancy varied according to lifestyle and habitat. Less variability was observed in the muscle profiles, characterized by low lipid content (<1.0% ww) and high relative levels of polar lipids (>70%). The lipid and fatty acid profiles of the kidney and pancreas showed the highest intraspecific variability, suggesting these organs also have complex roles in lipid storage and metabolism. Overall intra- and interspecific differences in the tissue fatty acid profiles could be related to differences in a number of factors including phylogeny, habitat (depth), buoyancy regulation and diet and presumably also reflect different ecological roles. The lipid and fatty acid profiles are the first published for Rhinochimaera pacifica, Chimaera lignaria and Figaro boardmani and the first to demonstrate interspecific variation in lipid profiles of various tissues of deepwater chondrichthyans. The application of multivariate analysis to lipid class and fatty acid tissue profiles in chondrichthyans inferred dietary differences and metabolic preferences between species and habitats. These results have important implications for the future use of fatty acids as dietary tracers in chondrichthyan research.     

Application of random amplified polymorphic DNA (RAPD) markers to evaluate intraspecific genetic variation in red mullet (Mullus barbatus)

The random amplified polymorphic DNA (RAPD) technique was used to evaluate genetic affinities among eight red mullet (Mullus barbatus L., 1758) samples from the Mediterranean Sea. Twenty-nine random primers were used. Despite the variability which was found within samples, no specific RAPD marker for the discrimination of the populations was detected. The data analysis revealed that the genetic diversity among populations is positively related to their geographic distances. The results of this study indicate that the estimated intraspecific variation may be more␣pronounced with RAPD markers than with allozymes when the two approaches are applied on the same populations. Received: 7 November 1997 / Accepted: 30 May 1998