Cryptic ecological diversification of a planktonic estuarine copepod, Acartia tonsa

The recent discovery of cryptic species in marine holoplankton, organisms that 'drift' in oceanic currents throughout their life cycle, contrasts with their potential for long-distance passive dispersal and presumably high gene flow. These observations suggest that holoplankton species are adapting to surprisingly small-scale oceanographic features and imply either limited dispersal or strong selection gradients. Acartia tonsa is a widespread and numerically dominant estuarine copepod containing deep mitochondrial lineages within and among populations along the northwestern Atlantic coast. In this study, we intensively investigated A. tonsa populations in Chesapeake Bay with the goals of testing species status for the deep lineages and testing for their association with environmental features over space and time. Phylogenetic analyses of DNA sequences from mitochondrial cytochrome c oxidase I (mtCOI) and the nuclear ribosomal internal transcribed spacer (nITS) resolved two concordant monophyletic clades. Deep divergence between the two clades (13.7% uncorrected sequence divergence for mtCOI and 32.2% for nITS) and genealogical concordance within sympatric populations strongly suggest that the two clades represent reproductively isolated cryptic species. Based on restriction fragment length polymorphisms of mtCOI, representatives from the two clades were found consistently associated with contrasting salinity regimes (oligohaline vs. meso-polyhaline) with an overlap between 2 and 12 PSU in samples from 1995 to 2005. Finding these patterns in one of the best-known estuarine copepods reinforces the conclusion that marine biodiversity is underestimated, not only in terms of species numbers, but also with respect to niche partitioning and the potential importance of ecological divergence in marine holoplankton.     

Feeding behaviour and migrations in a natural population of the copepod Acartia tonsa

This paper deals with the variations on feeding activities and diel migrations of Acartia tonsa Dana, the dominant copepod species in Berre lagoon (west Mediterranean French coasts).A 27 hour in situ study was carried out during June 1989, at a station located in the south west of the lagoon. Vertical profiles of salinity, temperature and dissolved oxygen taken each 12 hours showed a stratification of the water column in two distinct layers: (1) a superficial layer with higher temperature, moderate salinity, and high oxygen concentrations; (2) a colder, more saline and almost anoxic deep layer. In situ chlorophyll a measurements were made at –1, –3 and –5 m; concentrations were relatively homogeneous through the water column during the whole sampling period.Zooplankton samples were collected every 3 hours with a 200 µm mesh net, in three strata (0–2 m, 2–4 m, 4–6 m). A complete dominance of A. tonsa was observed in the zooplankton community. Our results point out clearly a nocturnal migration, with individuals concentrating in both superficial layers; thus an unimodal migratory pattern can be inferred. Gut flourescence measured following the Mackas & Bohrer technique (1976), showed higher values during night time, and values for females were the highest with wider day—night variations. Similar results were found in laboratory experiments with copepods fed with a culture of Dunaliella tertiolecta.Gut evacuation rate was measured in two laboratory experiments either mixing or separating males and females. Evacuation rate was 17.92 and 27.25 min for males and females respectively.Phytoplankton daily ration for A. tonsa calculated by gut flourescence and gut evacuation rate was particularly low, for it represents only 10% of the individual carbon weight.Moreover, grazing impact on phytoplankton is very restricted, it represents less than 1% of the daily phytoplankton stock.     

Cryptic diversity and comparative phylogeography of the estuarine copepod Acartia tonsa on the US Atlantic coast

Unexpectedly strong geographic structures in many cosmopolitan species of marine holoplankton challenge the traditional view of their unrestrained dispersal and presumably high gene flow. We investigated cryptic lineage diversity and comparative phylogeography of a common estuarine copepod, Acartia tonsa, on the US Atlantic coast, using mitochondrial (mtCOI) and nuclear (nITS) gene markers. Three broadly sympatric lineages (F, S, X) were defined by genealogically concordant clades across both gene trees, strongly supporting recognition as reproductively isolated species. Limited dispersal seems to have had a major role in population differentiation of A. tonsa in general, with gene flow propensities rank ordered X > S > F. Geographic structure was found only at large scales (1000–2000 km) in X and S. Phylogeographic patterns in all three lineages were mostly concordant with previously recognized zoogeographic provinces but a large mid‐Atlantic gap in the occurrence of lineage X, coupled with its presence in Europe, suggests possible nonindigenous origins. For lineage F, physiological adaptation to low‐salinity environments is likely to have accentuated barriers to gene flow and allopatric differentiation at both regional and continental scales. Three allopatric F sublineages inferred a southern centre of origin and a stepwise northward diversification history at the continental scale. The most recently derived F sublineages, in the mid‐Atlantic Bight, showed strong phylogeographic patterns at nITS albeit weaker at mtCOI. Applying a crustacean mtCOI molecular clock suggests that A. tonsa lineages diverged pre‐Pleistocene but mid‐Atlantic F lineage diversification may be post‐Pleistocene.     

Morphological and genetic differences in ecologically distinct populations of Petrosia (Porifera, Demospongiae)

The taxonomic status of two sponge tnorphotypes living sympatrically in Mediterranean caves and usually ascribed to Petrosia ficiformis was elucidated on the basis of morphological, morphometric and genetic features. The two morphotypes, spherical and cylindrical, showed differences in the shape and size of spicules and in the morphology of the aquiferous system. Electrophoretic analyses demonstrated that the two morphotypes are reproductively independent and so they should be considered as distinct biological species. The spicular features allow attribution of the cylindrical morphotype to P.ficiformis and the spherical one to P. clavata. For these two species parasympatric speciation is proposed.     

Seasonal variability in the individual carbon content of the calanoid copepod Acartia bifilosa from the northern Baltic Sea

The seasonal variation in the carbon content of the calanoid copepod Acartia bifilosa was studied in the northern Baltic Sea. Monthly length-weight relationships were followed from November 1990 to October 1991 by analysing the organic carbon content of individuals, using a high temperature combustion method. The monthly length (L) on carbon (C) regressions of copepodites were best described by power functions (C = aL^b), where 75% to 88% of the variation could be explained by length. Nauplii length explained less of their carbon content (63–71 %). The carbon content per length was highest in summer when the nutritional situation was good and temperature at its highest. However, the carbon-length regressions differed significantly between months, except in winter, when two subsequent months had similar regressions. The results pointed out the importance of seasonal variation as well as the risk of making errors if biomass estimates based on length on carbon regressions are used. Carbon analysis should, if possible, be done on every sample or should at least be tested as to whether the regressions are consistent with the study material before estimations are calculated.     

Morphological aspects of subitaneous and resting eggs from Acartia josephinae (Calanoida)

A. josephinae collected along the Salento Peninsula (Ionian and Adriatic Seas) laid two types of eggs, under laboratory conditions, and they are described for the first time. Under a light microscope subitaneous eggs looked smooth on the surface, but scanning electron microscopy revealed that the surface was not perfectly smooth but tubercular with many small protuberances. Resting eggs had long and thick spines whose apical part was multi-branched. The use of egg morphology as a taxonomic character, in the systematic of copepods, is proposed.     

Spatial variability of stable carbon and nitrogen isotope ratios in a Mediterranean coastal lagoon

Exploring the trophic pathway of organic matter within the Mauguio lagoon (southern France, western Mediterranean), we found spatial differences in the isotopic composition (both δ¹³C and δ¹⁵N values) of organic matter sources (primary producers, particulate and sedimentary organic matter), which were mirrored in the upper trophic levels (invertebrates and fish). On average, δ¹³C was heavier by about 1.5–2‰ in the location under marine influence than in the sites influenced by freshwater discharge. The opposite trend was found for δ¹⁵N, which attained maximum values in the north-central zone influenced by freshwater delivery. For both C and N stable isotope ratios, the highest spatial variability was found in organic matter sources (2–3‰), while invertebrates and fish exhibited less variability (\~1–2‰). The differences observed may be related to both anthropogenic (wastewater input) and natural (marine vs. terrestrial inputs) factors. Discharge of wastewater, which affects the innermost location, generally determines an increase in the relative abundance of ¹⁵N. In addition, terrestrially derived nutrients and organic matter, which also affect the innermost location, are known to determine a shift towards ¹³C-depleted values. Our results substantiate the finding that the analysis of carbon and nitrogen stable isotopes can help in elucidating origin and fate of organic matter in coastal lagoons, which are characterised by a great spatial variability and complexity.     

Morphological variation in the catadromous Australian bass, from seven geographically distinct riverine drainages

The Australian bass Macquaria novemaculeata is a catadromous species restricted to coastal drainages of south-eastern Australia and recently has undergone reductions in population numbers. Morphometric variation among bass from seven distinct drainages was used to test for the presence of stock structuring. Males and females were treated separately. For both male and female bass, significant heterogeneity in morphology among sample locales was revealed by univariate statistics and discriminant function analysis, with morphological variation found to differ in a clinal pattern. The demonstration of population-specific morphological traits among bass from different river drainages may indicate that inter-populational movement is restricted and that each sample locale constitutes a separate stock of bass.     

Temporal variability of the ichthyofauna in a Northern Aegean coastal lagoon (Greece). Influence of environmental factors

The ichtyofauna of the Porto-Lagos Lagoon in northern Greece was sampled over 17 months, using a bag-seine net, to investigate factors influencing species seasonal composition and abundance. Of the 35 resident, migrant and straggler species caught, representing 22 families, the most numerically dominant species were Atherina boyeri (46.4% of total catch), which today is the most economically important species, followed by Pomatoschistus marmoratus (28% of total catch) and Aphanius fasciatus (9.1% of total catch). Mugilidae and Sparidae were the most diverse families. According to various diversity indices, the lagoon’s ichthyofauna was found to be most diverse in May 1990; taxonomic spread however, as shown by the taxonomic distinctness index (Δ*), was lowest in July 1990, when the greatest monthly water temperature range was observed (13.5 °C). Overall fish relative abundance was positively correlated with the maximum monthly water temperature (r²=0.32, p < 0.05). The NMDS and cluster analysis performed on the bag seine catches showed four groupings, based mainly on the life history of the species. The main group consisted of all the resident species of the lagoon and four mugilid species that were present throughout most of the sampling period. The second group consisted of marine migrant species that spent a short period of their life in the lagoon. Marine migrants and one marine straggler that were found once or twice formed the third group, while the fourth group consisted of species with casual presence. The importance of Porto-Lagos Lagoon as nursery and feeding area for marine species was confirmed by the catch of young individuals of several marine species.     

Social and genetic characteristics of geographically isolated populations in the ant Formica cinerea

The ant Formica cinerea in northern Europe has geographically isolated populations that were examined using five microsatellite loci. The populations differ widely regarding the social organization of colonies. Based on genetic relatedness (r) among worker nest mates, the populations were classified as M type with monogynous (single queen) colonies (r > 0.59), as P type with polygynous colonial networks (r < 0.1), or as intermediate with weakly polygynous colonies (0.16 < r < 0.47). The social types showed weak geographical clustering, but the overall distribution indicated that the shift between the social types has occurred several times. The geographically isolated populations had slightly reduced levels of genetic diversity compared to populations from areas where the species is abundant and continuously distributed. Many of the isolated populations consisted of monogynous or weakly polygynous colonies, making their effective population sizes small, and some of them also showed weak bottleneck effects. The overall level of microsatellite diversity within populations was relatively high and differentiation among populations low, indicating recent connections. Isolation of populations may thus be a new phenomenon resulting from reduction of suitable habitats. At the local level, we obtained limited support from a group of nearby subpopulations in southern Finland to the hypothesis that the P type is connected to restricted dispersal. Other P type populations did not, however, show similar elevated levels of differentiation.     

Seasonal variability of copepod abundance in the Balearic region (Western Mediterranean) as an indicator of basin scale hydrological changes

Spatial and temporal changes of the copepod community have been investigated and related to the environmental variability of the Balearic Sea (Western Mediterranean). The period studied spans from 1994 to 1999 during which we analyzed the abundance and structure copepod variability over a cross-shore transect. Results showed a close link between hydrological changes and the variations of copepod abundance. The synchronous variability of copepods and hydrography indicated the rapid response of this zooplankton group to the inflow of cold and warm water masses coming through the study area. Cluster analysis revealed four main copepod assemblages that distinguished the coastal from the oceanic species and those species with different water masses preference. The copepod assemblage composed of Calanus helgolandicus, Clausocalanus arcuicornis, C. pergens, C. paululus, Calocalanus tenuis and Pleuromamma gracilis was associated with cool salty waters, whereas the assemblage formed by Temora stylifera, C. pavo, C. styliremis, Centropages bradyi and Acartia danae was related to warmer less saline Mediterranean waters. Moreover, it is suggested that changes in sea water temperature and salinity are linked to large-scale changes likely occurring at a basin scale, which is reflected in the Western Mediterranean mesoscale hydrographic changes. Therefore, it is stressed that changes in the Balearic copepod community can be used as potential tracers of the western Mediterranean water masses. Guest editors: S. Souissi & G. A. Boxshall Copepoda in the Mediterranean: Papers from the 9th International Conference on Copepoda, Hammamet, Tunisia     

Effects of food on bacterial community composition associated with the copepod Acartia tonsa Dana

The estuarine copepod Acartia tonsa naturally carried diverse strains of bacteria on its body. The bacterial community composition (BCC) remained very conservative even when the copepod was fed different axenic algal species, indicating that the food per se did not much affect BCC associated with the copepod. In xenic algal treatments, however, copepod-associated BCC differed with each alga fed, even though the same bacterial source was used to inoculate the algae. In addition, starved copepods taken at the same location but at different times significantly differed in their BCC. Algal species composition and copepod life history therefore serve to regulate BCC associated with copepods, and spatial and temporal variations in algal species composition and copepod origin would alter bacteria–copepod interactions.     

Waterbird communities along the estuarine salinity gradient of the Schelde estuary, NW-Europe

The zonation of non-breeding waterbirds along the Schelde estuary (The Netherlands–Belgium), one of the longest estuaries in NW-Europe with still a complete salinity gradient, including a large freshwater tidal area, was described. Numbers of birds were counted monthly over the period October 1991 to June 1997. Highest numbers of waterbirds were observed in late autumn and winter, with annual peak numbers ranging between 150,000 and 235,000 individuals for the whole estuary. Based on a multivariate analysis different waterbird communities were observed along the salinity gradient. The polyhaline areas of the estuary were numerically dominated by the waders Oystercatcher and Dunlin. Due to the presence of a large brackish marsh in the mesohaline zone, the waterbird community in this area was dominated by the herbivores Wigeon and Greylag Goose. In the oligohaline and freshwater tidal areas, the waterbird community was dominated by duck species, with Teal and Mallard being the most important. The international importance of the Schelde estuary for waterbirds was evidenced by the fact that for 21 waterbird species the 1% level criterion, according to the Ramsar convention, was exceeded. The relation of the observed diversity and community patterns with the functional and habitat diversity of the Schelde estuary as well as the effect of recent conservation measures to preserve this habitat were discussed.     

Albinism in phylogenetically and geographically distinct populations of Astyanax cavefish arises through the same loss-of-function Oca2 allele

The Mexican tetra, Astyanax mexicanus, comprises 29 populations of cave-adapted fish distributed across a vast karst region in northeastern Mexico. These populations have a complex evolutionary history, having descended from ‘old'' and ‘young'' ancestral surface-dwelling stocks that invaded the region ∼6.7 and ∼2.8 MYa, respectively. This study investigates a set of captive, pigmented Astyanax cavefish collected from the Micos cave locality in 1970, in which albinism appeared over the past two decades. We combined novel coloration analyses, coding sequence comparisons and mRNA expression level studies to investigate the origin of albinism in captive-bred Micos cavefish. We discovered that albino Micos cavefish harbor two copies of a loss-of-function ocular and cutaneous albinism type II (Oca2) allele previously identified in the geographically distant Pachón cave population. This result suggests that phylogenetically young Micos cavefish and phylogenetically old Pachón cave fish inherited this Oca2 allele from the ancestral surface-dwelling taxon. This likely resulted from the presence of the loss-of-function Oca2 haplotype in the ‘young'' ancestral surface-dwelling stock that colonized the Micos cave and also introgressed into the ancient Pachón cave population. The appearance of albinism in captive Micos cavefish, caused by the same loss-of-function allele present in Pachón cavefish, implies that geographically and phylogenetically distinct cave populations can evolve the same troglomorphic phenotype from standing genetic variation present in the ancestral taxon.     

Morphological and morphometrical analysis of Heterodera spp. populations in Jordan

Phenotypic diversity of five Jordanian populations of cyst nematodes, Heterodera spp. collected from five regions from Jordan (Ar-Ramtha, Madaba, Dana, Al-Karak, and Jerash) was investigated. Soil samples were collected from one representative field in each region. Morphological and morphometrical characteristics revealed that Heterodera latipons is dominated in cereal fields at Ar-Ramtha, Madaba, Dana and Al-Karak regions and Heterodera schachtii in Jerash. Cysts populations from all cereal fields had bifenestrate vulval cone and a strong underbridge. Wherever, cysts of the cabbage population had ambifenestrate vulval cone with long vulval slit. The bullae were absent in Ar-Ramtha, Madaba and Dana populations, but present in Al-Karak and Jerash. Based on 12 morphometrical characters, the first three functions in canonical discriminant analysis accounted 99.3% of the total variation. Distance from dorsal gland duct opening to stylet base, underbridge length, a = L/W (body length/midbody width) and length of hyaline tail tip had strong and significant contributions in the first function. While the second function was strongly influenced by length of hyaline tail, fenestral length, fenestral width and tail length. However, the third canonical discriminate function was found to be influenced by stylet length, fenestral length, a = L/W (body length/midbody width) and underbridge width. The graphical representation of the distribution of the samples showed that the first canonical discriminant function clearly separated H. schachtii from Jerash from other populations. Whereas, H. latipons collected from Madaba and Dana were clearly separated in the second function. The results indicated that differences at morphological and morphometrical levels revealed diverse populations of Heterodera spp. in Jordan.     

Induction of embryonic dormancy in the calanoid copepod Acartia hudsonica: heritability and phenotypic plasticity in two geographically separated populations

The planktonic copepod Acartia hudsonica apparently requires summer diapause or dormancy to persist in Narragansett Bay, RI. In estuaries to the north, however, active A. hudsonica populations are present year-round and may not express dormancy. Using a full-sibling rearing design and analysis of variance (ANOVA), I assessed the importance of phenotypic and genetic sources of variation in two geographically separate populations, one from Maine and one from Rhode Island. Both populations showed phenotypic plasticity in the percentage of dormant eggs produced. Moreover, experiments revealed significant sibship-environment interactions in both populations, indicating that the phenotypic plasticity has a genetic component. Both populations also revealed a significant amount of genetic variation in the percentage of dormant eggs produced. For the Maine population, broad sense heritability was high (0.91, 1.10) in two short-day (12L:12D) treatments and near zero (0.08) in the long-day treatment (15L:9D). For a Rhode Island population, broad sense heritability was higher (0.95) in a low-temperature short-day treatment (13.5 °C, 12L:12D) than in a high-temperature short-day treatment (17.5 °C, 12L:12D; h²=0.25).     

Genetic variability and structure of the remnant natural populations of <Emphasis Type="Italic">Cedrus libani</Emphasis> (Pinaceae) of Lebanon

Cedrus libani of Lebanon is a valuable natural resource and the dominant species in its natural ecosystem. Intense and diverse anthropogenic pressures over historical times raised concerns about its genetic vigor and continued survival. Our investigation of the genetic diversity included samples from all remnant natural populations. Assessment of the genetic diversity using random amplified polymorphic DNA markers revealed the persistence of considerable variation distributed within populations with low population differentiation corroborated by Bayesian and analysis of molecular variance estimates (G _ST = 0.07, Φ _ST = 0.09). Individual assignment tests were carried out to investigate measures of gene flow. Inferences concluded that this natural heritage is not currently threatened by inbreeding or by random genetic drift. Correlation studies investigated possible effects of spatial distribution and environmental conditions on genetic structure. A climatic trend corresponding to a temperature–humidity gradient correlated significantly with the level of genetic diversity, while the edaphic variation did not.