Observations on the embryonic development of some deep-living decapod crustaceans,with particular reference to species of Acanthephyra

Eggs of meso- and bathypelagic decapod crustaceans were removed from gravid females and allowed to develop in vitro. The time course of embryonic development, as indicated by the appearance of certain morphological and physiological features, was determined for most species at a constant temperature of 12°C. The embryonic period, and the time between the appearance of certain key features such as naupliar and compound eyes and heartbeats, showed little variation in any one species. Significant differences were observed in the development times of different species. In general, species with large eggs have a longer embryonic period than those with small eggs, but there also variations between species with eggs of similar size. During the development of Acanthephyra eggs there is an approximate doubling of the egg volume, correlated with a decrease in egg density and an increase in the water content. The increase in the water content and egg volume is similar to that of other decapod eggs. It is concluded that those species with small eggs could have several broods in 1 year, whereas those species with large eggs are likely to be restricted to one, or in some cases possibly two, broods.     

Field identification of ‘types’ A and B of the ascidian Ciona intestinalis in a region of sympatry

The ascidian species Ciona intestinalis is a major model chordate in developmental and evolutionary biology, and an important fouling organism and invasive species. However, genomic investigation has recently revealed the existence of two cryptic species, genetically distinct yet without obvious morphological differences, currently referred to as types A and B. Here, we show that they are externally distinctive in a zone of sympatry in the western English Channel. Examining genotyped specimens, we found that types A and B of C. intestinalis can generally be distinguished by body colour, pigmentation at the distal end of the siphons and the presence or absence of tubercles on the sides of the siphons. Detecting specimens of hybrid descent still requires detailed molecular analysis, but these visual characters in combination will identify living specimens of types A and B with high probability. These differences are shown to be inherited.     

Comparaison biochimique de quatre espèces de copépodes planctoniques de Méditerranée nord-occidentale à partir des enzymogrammes des estérases,de la leucine aminopeptidase,de la phosphatase alcaline,de la malate déshydrogénase et de l'enzyme malique

Four copepod species (Acartia clausi, Centropages typicus, Temora stylifera, Calanus helgolandicus) from the northwest Mediterranean were subjected to electrophoresis on polyacrylamide gel in an attempt to distinguish specific biochemical characteristics for each species. Enzymograms of 5 enzymes were studied: esterases, leucine aminopeptidase, alkaline phosphatase, malate dehydrogenase and the malic enzyme. Marked interspecific differences were revealed, enabling specific biochemical identities to be established which can be useful as taxonomic criteria in the same way as morphological characteristics. The study was performed during different seasons in 1980.     

Developmental and genetic evidence for the existence of three morphologically cryptic species of<Emphasis Type="Italic"> Crepidula</Emphasis> in northern Chile

The taxonomy of Crepidula has been based principally on shell morphology, but shell characteristics in this genus may be strongly influenced by the substrates they inhabit and are thus of limited use in classification. Four species in this genus are currently recognized from the Chilean coast, of which two, Crepidula dilatata and C. fecunda, are sympatric. These species are morphologically cryptic and differ only in their larval development: C. dilatata has indirect development with a planktonic phase, and C. fecunda direct development within egg capsules. The other two species, C. coquimbensis and C. philippiana, are species that occur within gastropod shells inhabited by hermit crabs and differ from the other species in internal morphology of the shell, developmental features, and geographic distribution. Analyses of nine morphological characters, eight developmental characters, and 26 enzymatic loci were carried out on four Crepidula populations in the Coquimbo region of northern Chile. Four populations of Crepidula were identified based on morphological, developmental, and protein electrophoresis features. Our results suggest the presence of C. coquimbensis plus three morphologically cryptic species including C. dilatata, C. fecunda, and a new previously unrecognized cryptic species. All four populations showed differences in larval development pattern. C. fecunda have indirect development, with free-swimming larvae. The remaining three species have intracapsular metamorphosis, but different modes of larval nutrition. C. dilatata have nurse eggs that are consumed by mechanical destruction; C. coquimbensis engulf nutritional embryos; and the new species eat nutritive embryos by rotation. Protein electrophoresis data support the specific condition of each group with different larval development and endorse the use of developmental differences in taxonomy of Crepidula. Genetic distance shows that C. dilatata and C. fecunda are closely related, and that C. coquimbensis is closer to them than to the new cryptic species.     

Life-history evidence for sibling species in Axiothella rubrocincta (Polychaeta: Maldanidae)

Individual Axiothella rubrocincta Johnson on the Californian coast live in U-shaped tubes, feed on the surface of the sediment, attain densities of 100 m^-2, attain lengths of 140 mm, have maximum oocyte diameters of 220 μm and produce demersal larvae. Individuals from Washington populations live in vertical tubes, feed 10–15 cm below the sediment surface, reach densities of 5000 m^-2, attain lengths of 60 mm, have maximum oocyte diameters of 385 μm and brood their young in their tubes to at least the 7-setiger stage. Individuals from California have a scalloped posterior margin of the cephalic plate, while individuals from Washington have entire margins. There are no other consistent morphological differences between populations. The population from Eagle Cove, Washington has individuals similar in size and oocyte diameter to individuals from California, but which maintain vertical tubes. A. rubrocincta represents a sibling species complex.     

Quaternary geographical sibling speciation and population structuring in the Eastern Atlantic skates (suborder Rajoidea) Raja clavata and R. straeleni

The European Raja clavata and the South African R. straeleni are marine skates which exhibit highly conserved morphological and ecological traits. Owing to this, taxonomic and evolutionary relationships between the two taxa have not yet fully elucidated. Here, we have tested the hypothesis that restricted gene flow and genetic divergence between these taxa might be associated with climatic/oceanographic discontinuities by surveying genetic variation in ten geographical samples at control region (CR) and amplified fragment length polymorphism (AFLP) loci. The clustering of CR haplotypes in two reciprocally monophyletic clades consistent with taxon zoogeography and the significant AFLP F values between the European and South African populations indicated the two taxa as recently diverged peripatric sibling species. Within each species, significant spatial genetic heterogeneity among samples at both markers revealed population structuring. We argued that structured populations and isolated sibling species might represent two stages of geographical speciation.     

Shell-boring versus tube-dwelling: is the mode of life fixed or flexible? Two cases in spionid polychaetes (Annelida, Spionidae)

Free crawling was an original life style, and the ability to build tubes and bore into shells evolved independently in various annelids and more than once within the family Spionidae. Absence of morphological differences between the shell-boring (SB) and tube-dwelling (TD) spionid worms and the innate ability of borers to build tubes raised a question whether the mode of life is fixed or flexible. Sequence data of three gene fragments of the mitochondrial 16S rDNA and nuclear 18S and 28S rDNA (in total 1,677 bp) have shown that SB and TD individuals of Dipolydora carunculata from the Sea of Japan were genetically identical, whereas those of Polydora triglanda from Taiwan were genetically distant and not conspecific. These data indicate that some species are constrained to a certain mode of life, while others are flexible and individuals can be either SB or TD depending on the place of settlement in the end of their larval development.     

A panmictic fiddler crab from the coast of Brazil? Impact of divergent ocean currents and larval dispersal potential on genetic and morphological variation in Uca maracoani

Marine species tend to exhibit relatively less population structuring than terrestrial species owing to fewer barriers to gene flow and increased connectivity resulting from greater dispersal abilities. Thus, in many cases, life history plays a more important role in phylogeography of marine taxa than do oceanographic features. Littoral species are of particular phylogeographic interest because they possess life histories that have both marine and terrestrial characteristics. This study evaluates the synergistic impact of divergent ocean currents and a high larval dispersal potential on the phylogeography of the fiddler crab, Uca maracoani, distributed along the coast of Brazil. Patterns of genetic variation were assessed with sequence data for a portion of the mitochondrial COI gene and AFLPs. Geometric morphometric techniques were used to evaluate morphological variation. Results revealed a lack of discernible genetic subdivision. However, geometric morphometrics showed statistically significant morphological differentiation. The absence of a clear phylogeographic pattern appears to be determined primarily by life history characteristics permitting a high level of connectivity. One, or a combination of several factors, may explain the incongruity between genetic and morphologic signatures, including phenotypic plasticity, incomplete lineage sorting, or recent and ongoing genetic divergence.     

Eurythoe complanata (Polychaeta: Amphinomidae), the ‘cosmopolitan’ fireworm, consists of at least three cryptic species

Eurythoe complanata (Pallas 1766) has been considered a cosmopolitan species with a great morphological similarity across its geographic range. To elucidate whether E. complanata is actually a single species, genetic (cytochrome oxidase subunit I and allozymes) and morphological differences were compared among specimens from the Pacific, Caribbean, and South Atlantic Oceans. Large levels of COI divergence (10–22%) and diagnostic allozyme loci identified three cryptic species: one in the eastern Pacific and two in the Atlantic, with one being morphologically differentiated and found only in islands. COI sequences between Pacific and Atlantic lineages were much more divergent than those of other transisthmian invertebrates, indicating their split before the Panama Isthmus closure or a faster evolutionary rate of COI for this species. The existence of two Atlantic species may be a consequence of parapatric speciation followed by a secondary invasion or even a sympatric speciation in the Atlantic oceanic islands.     

A molecular method for the identification of resting eggs of acartiid copepods in the Thau lagoon, France

Acartia and Paracartia species, often known to co-occur, can exhibit complex life cycles, including the production of resting eggs. Studying and understanding their population dynamics is hindered by the inability to identify eggs and early developmental stages using morphological techniques. We have developed a simple molecular technique to distinguish between the three species of the Acartiidae family (Acartia clausi, A. discaudata and Paracartia grani) that co-occur in the Thau lagoon (43°25′N; 03°40′E) in southern France. Direct amplification of a partial region of the mitochondrial cytochrome oxidase I gene by polymerase chain reaction and subsequent restriction fragment length polymorphism results in a unique restriction profile for each species. The technique is capable of determining the identity of individual eggs, including resting eggs retrieved from sediment samples, illustrating its application in facilitating population dynamic studies of this ubiquitous and important member of the zooplankton community.     

Live discrimination of Calanus glacialis and C. finmarchicus females: can we trust phenological differences?

Two key players in the Arctic and subarctic marine ecosystem are the calanoid copepods, Calanus finmarchicus and C. glacialis. Although morphologically very similar, these sibling species have different life cycles and roles in the Arctic pelagic marine ecosystem. Considering that the distribution of C. glacialis corresponds to Arctic water masses and C. finmarchicus to Atlantic water masses, the species are frequently used as climate indicators. Consequently, correct identification of the two species is essential if we want to understand climate-impacted changes on Calanus-dominated marine ecosystems such as the Arctic. Here, we present a novel morphological character (redness) to distinguish live females of C. glacialis and C. finmarchicus and compare it to morphological (prosome length) and genetic identification. The characters are tested on 300 live females of C. glacialis and C. finmarchicus from Disko Bay, western Greenland. Our analysis confirms that length cannot be used as a stand-alone criterion for separation. The results based on the new morphological character were verified genetically using a single mitochondrial marker (16S) and nuclear loci (six microsatellites and 12 InDels). The pigmentation criterion was also used on individuals (n = 89) from Young Sound fjord, northeast Greenland to determine whether the technique was viable in different geographical locations. Genetic markers based on mitochondrial and nuclear loci were corroborative in their identification of individuals and revealed no hybrids. Molecular identification confirmed that live females of the two species from Greenlandic waters, both East and West, can easily be separated by the red pigmentation of the antenna and somites of C. glacialis in contrast to the pale opaque antenna and somites of C. finmarchicus, confirming that the pigmentation criterion is valid for separation of the two species.     

Contribution à l'étude écologique de quelques zoanthaires tempérés et tropicaux

Within the framework of an ecological investigation on zoanthids, 3 genera and 6 species have been studied from the vicinity of Marseilles, and 5 genera including 11 species analysed from near Tulear, southwest coast of Madagascar. The tropical forms are mainly distributed throughout the intertidal zone. These zoanthids require vigorously moving waters and a high level of salinity. Sensibility to pollution differs according to species. Distribution is based upon the degree of dampness and upon the nature of the bottom. In intertidal zoanthids, the degree of desicoation determines the vertical zonation of species occurring on the reef. Light is an important distributional factor in species which are mostly submerged, especially when they are associated with symbiotic algal cells. Another major factor is the composition of the sediment, although this is less important for the tropical species studied (except for Isaurus spongiosus Andre, 1877). A model is proposed for comparison of species in terms of sensibility to environmental factors. In temperate areas, zoanthids appear to be “mixticoles”, but the formation of the incrustation present in many zoanthids is not connected with the composition of the substrate. From an ethological point of view, temperate zoanthids differ from tropical species. The differences manifest themselves in morphological features and in behaviour. Since the macrocnemic zoanthids mainly dwell in temperate waters and brachycnemic zoanthids in tropical waters, it may be possible to extend these differences to macrocnemic and brachycnemic groups.     

Morphological, reproductive, and genetic variability among three populations of Crucibulum quiriquinae (Gastropoda: Calyptraeidae) in northern Chile

Crucibulum quiriquinae (Lesson, 1830) is the only species of Crucibulum currently recognized in northern Chile. Recent analysis of three Crucibulum populations obtained in northern Chile demonstrates the existence of morphological, genetic, and reproductive differences among populations. Two populations present in Bahía Tongoy (30°15'S), one inhabiting the shells of the snail Turritella cingulata and the other shells of the pectinid Argopecten purpuratus, showed morphological differences. However, both had planktonic larval development and show low genetic divergence (D=0.002). A third population from Bahía La Herradura (29°58'S), which also inhabits the shells of T. cingulata, did not show morphological differences compared with its counterpart from Bahía Tongoy. However the Bahía La Herradura population had intracapsular development and metamorphosis, and a larger genetic distance (D=0.06) from both Tongoy populations. The results of the reproductive and genetic analyses strongly suggested that the two Tongoy populations, although showing morphological differences, are biologically the same species, Crucibulum quiriquinae, whereas the La Herradura population is a new species.     

Enzymatic and morphological criteria for distinguishing between Cardium edule and C. glaucum of the Portuguese coast

Cardium edule L. and C. glaucum Bruguière are two sibling species of cockles commonly found in littoral systems along the Portuguese coast, where they sometimes occur as mixed populations. The problem of morphological separation of these two cockle species has not been completely solved. Indeed, the distinctive morphological criterion proposed by Brock after identifying the specimens by means of electrophoretic patterns of hepatopancreatic malate dehydrogenase was not effective when applied to Portuguese Cardium spp. The object of the present study was to establish morphological discrimination criteria for distinguishing Portuguese C. edule and C. glaucum on the basis of initial identification of individuals through the enzymatic criterion established by Brock. Cockles were collected from 1988 to 1991 at 16 estuarine and lagoonal systems along the Portuguese coast. Hepatopancreatic extracts were submitted to horizontal starch gel electrophoresis following Brock's procedure. The malate dehydrogenase patterns obtained were identical to those found by Brock. Clear interspecific differences in the leucine aminopeptidase and esterase isozyme systems were also detected. The morphological study of the shells involved 7 morphometric and 9 qualitative and semiquantitative characters. Canonical variate analysis of the morphometric characters showed that these could not be used for identification purposes. However, a clear separation between the two species was obtained by a principal-component analysis of qualitative and semiquantitative characters. The characters mainly useful for distinguishing between the two species were valve profile, shape of ventral valve junction, type of calcareous scales and shell colour.     

Investigating stock structure and trophic relationships among island-associated dolphins in the oceanic waters of the North Atlantic using fatty acid and stable isotope analyses

Short-beaked common dolphins (Delphinus delphis) and Atlantic spotted dolphins (Stenella frontalis) are the two most abundant cetacean species in the oceanic waters of Madeira and the Azores. They are of similar size, occur in similar habitats and are regularly observed in mixed-species groups to forage together. Genetic analyses suggested that, within each species, dolphins ranging around both archipelagos belong to the same panmictic population. We tested the hypotheses that (1) within each species, individuals from the two archipelagos belong to a single ecological stock; (2) between species, common and spotted dolphins have distinct trophic niches; using fatty acid (FA) and stable isotope (SI) analyses. Fatty acids and stable isotopes were analysed from 86 blubber and 150 skin samples of free-ranging dolphins, respectively. Sex-related differences were not significant, except for common dolphin FA profiles. In S. frontalis, FA and SI differences between archipelagos suggested that individuals belonged to different ecological stocks, despite the existence of gene flow between the two archipelagos. In D. delphis, differences were more pronounced, but it was not possible to distinguish between stock structure and a seasonal effect, due to differential sampling periods in the Azores and Madeira. Inter-specific comparisons were restricted to the Azores where all samples were collected during summer. Differences in FA proportions, noticeably for FA of dietary origin, as well as in nitrogen SI profiles, confirmed that both species feed on distinct resources. This study emphasizes the need for an integrated approach including both genetic and biochemical analyses for stock assessment, especially in wide-ranging marine top predators.     

Convergent development of ecological,genetic, and morphological traits in native supercolonies of the red ant Myrmica rubra

Ant supercolonies (large networks of interconnected nests) represent the most extreme form of multi-queen breeding (polygyny) and have been found across ant lineages, usually in specific long-term stable populations. Many studies on the genetic population structure and demography of ant supercolonies have been done in recent decades, but they have lacked multicolonial control patches with separated colonies headed by a single or few queens so the origin of the supercolonial trait syndrome has remained enigmatic. Here, we set out to compare sympatric supercolonial and multicolonial patches in two natural Danish populations of the common red ant Myrmica rubra. We used DNA microsatellites to reconstruct genetic colony/population structure and obtained morphological and density measurements to estimate life history and ecology covariates. We found that supercolonies in both populations completely dominated their patches whereas colonies in multicolonial patches coexisted with other ant species. Supercolony patches had very low genetic differentiation between nests, negligible relatedness within nests, and lower inbreeding than multicolonial patches, but there were no significant morphological differences. One population also had nests that approached true outbred monogyny with larger workers and males but smaller queens than in the two other social nest types. Our results suggest that once smaller colonies start to adopt additional queens, they also gain the potential to ultimately become supercolonial when the habitat allows rapid expansion through nest budding. This is relevant for understanding obligate polygyny in ants and for appreciating how and why introduced North American populations of M. rubra have recently become invasive.     

Repeated geographic divergence in behavior: a case study employing phenotypic trajectory analyses

Environmental effects on behavior have long been a focus of behavioral ecologists. Among the important drivers of behavior is predation environment, which can include the presence/absence of predators, differences in resource availability, and variation in individual density. Environments with predators are often more ecologically complex and “risky” than those without predators. Populations from these environments are sometimes more active and explorative than populations from low-risk, less complex environments. To date, most comparative studies of behavior are limited to within-species comparisons of populations from divergent environments, but neglect comparisons between species following speciation, thus limiting our understanding of post-speciation behavioral evolution. Brachyrhaphis fishes provide an ideal system for studying correlations between divergent environments and behavior within and between species. Here, we test for differences in two behavioral traits—activity and exploration —between sister species Brachyrhaphis roseni and Brachyrhaphis terrabensis that occur in divergent predation environments. Species differed in activity and exploration, with higher activity and exploration levels in populations that co-occur with predators. Furthermore, we found drainage-by-species interactions, indicating that the nature of divergence varied geographically. Using the recently developed phenotypic trajectory analysis (PTA), we quantified this difference and found that, while the geographically isolated populations of sister species tended to evolve in parallel, the magnitude of divergence between species differed between drainages. Our results highlight the utility of PTA for multivariate behavioral data and corroborate past predictions that complex and risky environments are correlated with increased activity and exploration levels and that divergence continues post-speciation.     

Microbial activities as a function of water depth in the Ligurian Sea: an autoradiographic study

Fourteen species of sergestid shrimps were collected in the Sargasso Sea between the surface and 1500 m near Bermuda on 4 cruises. The vertical distribution and feeding activity of the most abundant species are discussed in relation to interspecific competition and the adaptive significance of vertical migration. Each species lives within a narrow depth range and exhibits a diel vertical migration. Sergestes splendens migrated as much as 825 m, while S. japonicus migrated less than 100 m. Neither the seasonal nor permanent thermocline influenced the migration range. The only species which occurred together both day and night were S. pectinatus with S. vigilax and S. pectinatus with S. sargassi. Morphological differences in the third maxillipeds of these species suggest differences in feeding. Although most species eat a variety of organisms, the foreguts of S. grandis, S. corniculum, and S. splendens contained euphausiids more often than those of other species, and S. grandis and S. robustus fed more frequently on fishes. In contrast, S. japonicus appears to feed on detritus. Food was found in the foreguts of most species less frequently during the day than night, but no species fed only at night. S. sargassi and S. pectinatus fed equally day and night.     

Differences in kelp morphology between wave sheltered and exposed localities: morphologically plastic or fixed traits?

The ability of algae to change the shape of their thallus in response to the environment may be of functional and ecological importance to the alga, with many species of macroalgae exhibiting a great range of morphological variation across wave exposure gradients. However, differences in morphology detected between sheltered and exposed environments cannot determine whether such differences represent plastic responses to the local environment or whether morphology is genetically fixed. This study tested for differences in the morphology of the common kelp, Ecklonia radiata, between wave sheltered and exposed environments, and reciprocally transplanted juveniles to distinguish the nature of such differences (i.e. plastic vs fixed traits). Differences between exposure environments were consistent with known effects of exposure (i.e. a wide, thin thallus at sheltered sites and a narrow, thick thallus with a thick stipe at exposed sites). The reciprocal transplant experiment confirmed that morphological plasticity was the mechanism enabling this alga to display different patterns in morphology between exposure environments. Individuals transplanted to the exposed environment underwent a rapid and extreme response in morphology, which was not apparent in individuals transplanted to the sheltered environment that responded more slowly. These results suggest that stressors typical of sheltered environments (i.e. diffusion stress) may not be as influential (if at all) compared to stressors typical of exposed environments (i.e. breakage, dislodgement) in differentiating morphological characters between exposure environments.     

How wave exposure, group size and habitat complexity influence foraging and population densities in fishes of the genus Halichoeres (Perciformes: Labridae) on tropical rocky shores

Wave exposure and habitat complexity have been used to explain variations in the distribution patterns and behavior of many reef fishes. This study analyzed the influence of both factors on density and foraging activity, and the influence of group size on foraging in three species of the genus Halichoeres (Labridae) on tropical rocky shores. It was shown that initial phases (IP) and terminal phases (TP) Halichoeres, were influenced by wave exposure, although foraging in Halichoeres brasiliensis TP was not influenced by wave exposure. The IP in all three species were positively associated with rugosity and algal cover while the TP was positively associated with depth. Habitat complexity also influences foraging in these species. Group size influenced foraging activity, except in H. brasiliensis TP. We also found variations in microhabitat patches used for foraging between species and differences in the stomach contents between species and phases, showing that functional roles change in response to ontogenetic shifts.