Historical colonization and demography of the Mediterranean damselfish, Chromis chromis

The desiccation of the Mediterranean Sea during the Messinian Salinity Crisis 6.0-5.3 million years ago (Ma), caused a major extinction of the marine ichthyofauna of the Mediterranean. This was followed by an abrupt replenishment of the Mediterranean from the Atlantic after the opening of the Strait of Gibraltar. In this study, we combined demographic and phylogeographic approaches using mitochondrial and nuclear DNA markers to test the alternative hypotheses of where (Atlantic or Mediterranean) and when (before or after the Messinian Salinity Crisis) speciation occurred in the Mediterranean damselfish, Chromis chromis. The closely related geminate transisthmian pair Chromis multilineata and Chromis atrilobata was used as a way of obtaining an internally calibrated molecular clock. We estimated C. chromis speciation timing both by determining the time of divergence between C. chromis and its Atlantic sister species Chromis limbata (0.93-3.26 Ma depending on the molecular marker used, e.g. 1.23-1.39 Ma for the control region), and by determining the time of coalescence for C. chromis based on mitochondrial control region sequences (0.14-0.21 Ma). The time of speciation of C. chromis was always posterior to the replenishment of the Mediterranean basin, after the Messinian Salinity Crisis. Within the Mediterranean, C. chromis population structure and demographic characteristics revealed a genetic break at the Peloponnese, Greece, with directional and eastbound gene flow between western and eastern groups. The eastern group was found to be more recent and with a faster growing population (coalescent time = 0.09-0.13 Ma, growth = 485.3) than the western group (coalescent time = 0.13-0.20 Ma, growth = 325.6). Our data thus suggested a western origin of C. chromis, most likely within the Mediterranean. Low sea water levels during the glacial periods, the hydrographic regime of the Mediterranean and dispersal restriction during the short pelagic larval phase of C. chromis (18-19 days) have probably played an important role in C. chromis historical colonization.     

Historical population dynamics and demography of the Eastern Atlantic pomacentrid Chromis limbata (Valenciennes, 1833)

Recent studies have focused on the relationship between the marine fauna of the Eastern Atlantic and the Mediterranean Sea, but within the Atlantic, little is known about genetic relationships between populations of the Macaronesian islands. In this study, we tested whether the paleo-climatology and paleo-oceanography of the region could predict the genetic relationships among three Eastern Atlantic populations (Azores, Madeira, and Canaries) of a damselfish, Chromis limbata, and compared our results with its Mediterranean and adjacent Atlantic sister species, Chromis chromis. We combined phylogeographic and coalescent approaches using the fast evolving mitochondrial control region gene. No population structure was found for the three archipelagos. The coalescence time estimated for C. limbata (0.857-1.17 Mya) was much greater than that estimated for C. chromis. We propose that this difference reflects differences in glaciating extents in the Northeastern Atlantic and the Mediterranean. Diversity indexes (Hd and genetic distances) together with historical demographic parameters of C. limbata (Theta and g) revealed a more stable population history when compared to C. chromis. Our results suggest that the Macaronesian populations of C. limbata have probably been less affected by the last glaciation than the Mediterranean populations of C. chromis. Migration across the three archipelagos was estimated and a prevailing northwest trend was detected. This result supports the idea of a colonization of the Azores by warm water fish from Madeira or the westernmost Canary islands which acted as major glacial refugia for the tropical and subtropical marine fauna during the glaciations.     

Reproductive synchrony in a temperate damselfish, <Emphasis Type="Italic">Chromis limbata</Emphasis>

Synchronous reproduction has been reported in many tropical damselfishes, and it appears that temperature is the main driver of synchrony at these latitudes. However, reproductive synchrony and its relation to temperature is less well understood in temperate damselfish. The endemic Azores chromis (Chromis limbata) was chosen as a study species to address this topic. First, an ethogram of the reproductive behaviour and embryo development of the Azores chromis is provided. Then, are investigated (1) the reproductive dynamics at the colonial and inter-colonial levels, (2) how these are affected by temperature, and (3) the relation between reproductive dynamics and the social organization of temperate damselfish. In the Azores chromis, reproduction was cyclic with a high periodicity of 7.2 ± 0.63 days on average. Embryo development was synchronous within colonies towards the beginning and the end of incubation, but not at intermediate stages of development. Colonies that were closer were more synchronized than those located further apart, despite contrasting temperature and depth profiles. Embryo size differences were also greater between colonies that were further apart, and embryos tended to grow larger in warmer water. A literature review revealed that reproductive synchrony was common in colonial breeding damselfish species. Although these results suggest that within the breeding season, the timing and length of reproductive cycles cannot be explained by temperature alone, the observed changes in embryo size raise concern about the effects of changing oceanic conditions on this species.     

Effects of nautical traffic and noise on foraging patterns of Mediterranean damselfish (Chromis chromis)

Chromis chromis is a key species in the Mediterranean marine coastal ecosystems where, in summer, recreational boating and its associated noise overlap. Anthropogenic noise could induce behavioural modifications in marine organisms, thereby affecting population dynamics. In the case of an important species for the ecosystem like C. chromis, this could rebound on the community structure. Here, we measured nautical traffic during the summer of 2007 in a Southern Mediterranean Marine Protected Area (MPA) and simultaneously the feeding behaviour of C. chromis was video-recorded, within both the no-take A-zone and the B-zone where recreational use is allowed. Feeding frequencies, escape reaction and school density were analysed. C. chromis specimens were also collected from 2007 to 2008 to evaluate their physiological state using the Body Condition Index as a proxy of feeding efficiency. The MPA was more exploited by nautical tourism during holidays than on weekdays, particularly in the middle of the day. Greater traffic volume corresponded with lower feeding frequencies. The escape reaction was longer in duration (>1 min) when boat passed nearby, while moored boats did not induce an escape response. We found no differences in density between schools in the A- and B-zones and worse body conditions among those individuals inhabiting the B-zone in one area only. Overall, our findings revealed a significant modification of the daily foraging habits of C. chromis due to boat noise, which was slightly buffered by no-take zones established within the MPA.     

Relationship between roving behaviour and the diet and client composition of the cleaner fish Labroides bicolor

Diet analyses and observations of cleaning behaviour of two cleaner fishes revealed that Labroides bicolor fed more on client mucus, but Labroides dimidiatus fed more on ectoparasites, and that L. bicolor interacted with fewer species (36 species) compared with L. dimidiatus (44 species). The client species which contributed most to the dissimilarity between cleaner species were the dusky farmerfish Stegastes nigricans and bicolor chromis Chromis margaritifer damselfishes, which L. dimidiatus interacted with more often than L. bicolor, and the striated Ctenochaetus striatus and brown Acanthurus nigrofuscus surgeonfishes, which L. bicolor interacted with more; L. bicolor interacted with all parrotfishes (Scaridae) more. These results confirm the importance of repeated interactions and partner choice in determining the nature of interactions in mutualisms.     

Westward range expansion of the Lessepsian migrant Fistularia commersonii (Fistulariidae) in the Mediterranean Sea, with notes on its parasites

One adult female bluespotted cornetfish Fistularia commersonii was captured near the coast of Arbatax (Sardinia, Italy) in late October 2005. Macroscopic and microscopic analysis of the gonad revealed a postspawning ovary. Several body parts of the damselfish Chromis chromis were found in its stomach contents. Different developmental stages of 58 metazoan parasites belonging to Arthropoda, Nematoda and Platyhelminthes were detected in its mouth and digestive tract. The lepocreadiid Allolepidapedon fistulariae was reported for the first time from the Mediterranean and, with the exception of this species, all the parasites found were reported for the first time in this host. This new record of F. commersonii may confirm the rapid range expansion of this Lessepsian migrant in the western Mediterranean Sea, as well as the presence of its endoparasite A. fistulariae .     

Multidimensional polarization sensitivity in damselfishes

Using electroretinogram recording and microspectrophotometry we investigated spectral sensitivity and ultraviolet polarization sensitivity in three species of coral reef fishes commonly known as damselfishes. Here we show that three species of damselfishes (three-spot damselfish, Dascyllus trimaculatus; blacktail damselfish, D. melanurus; and blue-green chromis, Chromis viridis) have four classes of cone photoreceptors (lambda(max) ranges: ultraviolet 357-367 nm; short wavelength-sensitive 469-478 nm; medium wavelength-sensitive 482-493 nm; long wavelength-sensitive 512-524 nm; rods 499-500 nm). The three species shared similar combined spectral sensitivity but surprisingly complicated and varied polarization sensitivity. Damselfish examined in this study have three and four channel polarization sensitivity, the most complex polarization sensitivity recorded for any vertebrate. Such capacity could play an important role in mediating a conspecific visual communication network utilizing polarized light signals in the coral reef environment.     

Ecological differences between hamlet (Hypoplectrus: Serranidae) colour morphs: between-morph variation in diet

Dietary differences between hamlet Hypoplectrus spp. colour morphs were examined in fishes from Puerto Rico, U.S. Virgin Islands, Curacao, Honduras and Belize. Hamlet diet across all countries was characterized by large overlap between most colour morphs in both the proportion and numbers of dietary items consumed, although some differences between morphs were apparent. Indigo hamlets Hypoplectrus indigo were the only morph to consume fishes (blue chromis Chromis cyanea and sunshinefish Chromis insolata ) almost exclusively. The sympatric occurrence of other ecologically indistinguishable colour morphs, however, suggests that divergent ecological selection alone cannot explain population divergence in hamlets. Geographical variation in diet was also observed within black Hypoplectrus nigricans and yellowtail Hypoplectrus chlorurus hamlets which may reflect geographical differences in prey availability or differences in prey choice.     

Host-dependent differences in measures of condition associated with <Emphasis Type="Italic">Anilocra</Emphasis> spp. parasitism in two coral reef fishes

Parasites account for over half of the biodiversity on coral reefs, yet their ecological impacts are poorly understood. Cymothoid isopods of the genus Anilocra are large, conspicuous ectoparasites of coral reef fishes. French grunt (Haemulon flavolineatum) and brown chromis (Chromis multilineata) are commonly infected by Anilocra spp. in the Caribbean. These fishes play a significant role in trophic connectivity through their foraging and activity patterns, and Anilocra spp. infection has been reported to influence the trophic interactions of some fishes. Yet, how these changes manifest physiologically has not been quantified. Thus to determine the energetic effects of Anilocra spp. on French grunt and brown chromis, the relationships between Anilocra spp. infection and condition factor, percent moisture in the muscle tissue, total muscle tissue calories, and gut content volume were examined. The results of these analyses revealed that A. haemuli-infected French grunt had greater percent moisture in the muscle tissue but similar condition scores, calorie values, and gut content volumes compared to uninfected conspecifics. By comparison, Anilocra chromis-infected brown chromis had reduced condition factor, but similar percent moisture in the muscle tissue and total muscle tissue calories, as compared to uninfected conspecifics. This study provides evidence that infection by parasites of the same genus and within the same localities can have differential effects on fish host species, such that generalizations about the effects of parasitism across and within genera should be made cautiously.     

The recent colonization of south Brazil by the Azores chromis Chromis limbata

The damselfish Chromis limbata is native to the Macaronesian Archipelagos (Azores, Madeira and Canaries) and the western coast of Africa between Senegal and Angola. During the austral summers of 2008 and 2009 the species was recorded for the first time in the south‐western Atlantic Ocean around Campeche and Xavier Islands, in Florianópolis, Santa Catarina State, Brazil. Here, the progression of C. limbata in southern Brazilian waters is described using visual counts and genetic surveys and changes in the density of the native congener Chromis multilineata were also investigated. Underwater visual censuses of both Chromis species were carried out from 2009 to 2014. Chromis limbata tissue samples were collected and the mtDNA control region was sequenced and compared with mtDNA haplotypes from the natural range to confirm species identity, compare genetic diversity and to infer connectivity between newly established Brazilian populations. The Brazilian population of C. limbata increased significantly over the past 5 years and the effect on C. multilineata is still an open question, longer time‐series data will be necessary to clarify possible interactions. The molecular analyses confirmed species identity, revealed strong haplotype connectivity among Brazilian study sites and showed a low genetic diversity in Brazil when compared with the native populations, suggesting few individuals started the invasion. Four hypotheses could explain this colonizing event: C. limbata was released by aquarium fish keepers; larvae or juveniles were transported via ship ballast water; the species has rafted alongside oil rigs; they crossed the Atlantic Ocean through normal larval dispersal or naturally rafting alongside drifting objects. The rafting hypotheses are favoured, but all four possibilities are plausible and could have happened in combination.     

Megalopae and early postlarval stages of East African Percnon (Decapoda: Brachyura: Grapsidae)

The megalopae and first crab instars are described for three of the four Indo-west Pacific species of the grapsid crab Percnon—P. abbreviatum, P. guinotae and P. planissimum . Keys are provided to these three species and for the Atlantic/East Pacific P. gibbesi . The validity of separating P. planissimum and P. gibbesi is discussed, as are the implications of the unusually large megalopa in Percnon .     

Fishing top predators indirectly affects condition and reproduction in a reef-fish community

To examine the indirect effects of fishing on energy allocation in non-target prey species, condition and reproductive potential were measured for five representative species (two-spot red snapper Lutjanus bohar, arc-eye hawkfish Paracirrhites arcatus, blackbar devil Plectroglyphidodon dickii, bicolour chromis Chromis margaritifer and whitecheek surgeonfish Acanthurus nigricans) from three reef-fish communities with different levels of fishing and predator abundance in the northern Line Islands, central Pacific Ocean. Predator abundance differed by five to seven-fold among islands, and despite no clear differences in prey abundance, differences in prey condition and reproductive potential among islands were found. Body condition (mean body mass adjusted for length) was consistently lower at sites with higher predator abundance for three of the four prey species. Mean liver mass (adjusted for total body mass), an indicator of energy reserves, was also lower at sites with higher predator abundance for three of the prey species and the predator. Trends in reproductive potential were less clear. Mean gonad mass (adjusted for total body mass) was high where predator abundance was high for only one of the three species in which it was measured. Evidence of consistently low prey body condition and energy reserves in a diverse suite of species at reefs with high predator abundance suggests that fishing may indirectly affect non-target prey-fish populations through changes in predation and predation risk.     

Halacaridae (Acari) von der Atlantikküste des borealen Nordamerikas

38 halacarid species from the coasts of the boreal West Atlantic Ocean have been recorded. In a wide-spread net of stations, samples were taken from different substrata in marine and brackish waters in order to obtain information on the biology and ecology of halacarid species. Several habitats with their flora, fauna and halacarid species are described. In intertidal areas on the coasts of boreal North America fewer species were found than known from European coasts. Similar habitats on the west and east coasts of the North Atlantic Ocean are compared with respect to their halacarid population. In the rhombognathine and the genusHalacarellus, most species found in the West Atlantic Ocean are known in the East Atlantic too, living in similar habitats. The generaAnomalohalacarus andCopidognathus are common both in North American and European waters, but comparable biotopes are inhabited by different species, though often related or very similar in their appearance. 45 % of the halacarid species found in the boreal West Atlantic Ocean are also known in the East Atlantic. Hypotheses as to the dispersal and geographical distribution of halacarid genera and species are discussed. It is supposed, that many of the amphiatlantic species invaded biotopes on the American and European coasts, shortly after these continental plates drifted apart.     

Diversity, structure and function of fish assemblages associated with Posidonia oceanica beds in an area of the eastern Mediterranean Sea and the role of non-indigenous species

Temporal and spatial variation in density, biomass and body size of littoral fish species associated with nearshore Posidonia oceanica meadows was studied over an annual cycle in an area of the eastern Mediterranean Sea. A total of 109,350 littoral fishes were collected, belonging to 34 families and 88 species. Density of fishes peaked during the summer due to high numbers of juveniles. Season was a significant factor determining density, although number of species and biomass did not show any obvious seasonal pattern. Throughout the study, schooling planktivorous fish species such as the picarel Spicara smaris, the bogue Boops boops and the damselfish Chromis chromis were dominant, both in terms of density (80%) and biomass (70%). Temporal variation in density and body size of fishes was used to assess the seasonal and ontogenetic habitat use of each species, with their affinity to seagrass assessed by comparing their respective distribution on sand. Four functional guilds were created (juvenile migrants, seagrass residents, seasonal migrants and occasional visitors) to describe the habitat use of P. oceanica meadows by each species. Several species associated with P. oceanica meadows used this habitat mainly as juveniles during summer, although many others were present concurrently as adults and as juveniles. Among the species encountered, 11 were non-indigenous of Indo-Pacific origin, of which three used seagrasses mainly as juveniles and four as residents. The non-indigenous silverstripe blaasop Lagocephalus sceleratus ranked among the 10 most dominant species in terms of biomass (2%) and was classified as a seagrass resident.     

Phylogeography of Ophioblennius: the role of ocean currents and geography in reef fish evolution

Many tropical reef fishes are divided into Atlantic and East Pacific taxa, placing similar species in two very different biogeographic regimes. The tropical Atlantic is a closed ocean basin with relatively stable currents, whereas the East Pacific is an open basin with unstable oceanic circulation. To assess how evolutionary processes are influenced by these differences in oceanography and geography, we analyze a 630-bp region of mitochondrial cytochrome b from 171 individuals in the blenniid genus Ophioblennius. Our results demonstrate deep genetic structuring in the Atlantic species, O. atlanticus, corresponding to recognized biogeographic provinces, with divergences of d = 5.2-12.7% among the Caribbean, Brazilian, St. Helena/Ascension Island, Gulf of Guinea, and Azores/Cape Verde regions. The Atlantic phylogeny is consistent with Pliocene dispersal from the western to eastern Atlantic, and the depth of these separations (along with prior morphological comparisons) may indicate previously unrecognized species. The eastern Pacific species, O. steindachneri, is characterized by markedly less structure than O. atlanticus, with shallow mitochondrial DNA lineages (dmax = 2.7%) and haplotype frequency shifts between locations in the Sea of Cortez, Pacific Panama, Clipperton Island, and the Galapagos Islands. No concordance between genetic structure and biogeographic provinces was found for O. steincdachneri. We attribute the phylogeographic pattern in O. atlanticus to dispersal during the reorganization of Atlantic circulation patterns that accompanied the shoaling of the Isthmus of Panama. The low degree of structure in the eastern Pacific is probably due to unstable circulation and linkage to the larger Pacific Ocean basin. The contrast in genetic signatures between Atlantic and eastern Pacific blennies demonstrates how differences in geology and oceanography have influenced evolutionary radiations within each region.     

Fickle or Faithful: The Roles of Host and Environmental Context in Determining Symbiont Composition in Two Bathymodioline Mussels

The Mediterranean Sea and adjoining East Atlantic Ocean host a diverse array of small-sized mussels that predominantly live on sunken, decomposing organic remains. At least two of these, Idas modiolaeformis and Idas simpsoni, are known to engage in gill-associated symbioses; however, the composition, diversity and variability of these symbioses with changing habitat and location is poorly defined. The current study presents bacterial symbiont assemblage data, derived from 454 pyrosequencing carried out on replicate specimens of these two host species, collected across seven sample sites found in three oceanographic regions in the Mediterranean and East Atlantic. The presence of several bacterial OTUs in both the Mediterranean Sea and eastern Atlantic suggests that similar symbiont candidates occur on both sides of the Strait of Gibraltar. The results reveal markedly different symbiotic modes in the two species. Idas modiolaeformis displays high symbiont diversity and flexibility, with strong variation in symbiont composition from the East Mediterranean to the East Atlantic. Idas simpsoni displays low symbiont diversity but high symbiont fidelity, with a single dominant OTU occurring in all specimens analysed. These differences are argued to be a function of the host species, where subtle differences in host evolution, life-history and behaviour could partially explain the observed patterns. The variability in symbiont compositions, particularly in Idas modiolaeformis, is thought to be a function of the nature, context and location of the habitat from which symbiont candidates are sourced.     

Effects of depth and bottom communities on the distribution of highly territorial reef fish in the northwestern region of Cuba

The main goal of the study was to investigate changes in reef fish species abundance and its correlation with selected environmental variables. Three main questions guided the analyses: (i) Do the analytical methods have a significant influence on the results; (ii) What are the main assemblage variation axes; and (iii) What are the factors correlating best with assemblage composition. Highly territorial fish assemblages of several coral reefs along the northwestern region of Cuba were assessed using a stationary point visual census technique. A total of 39 701 individuals of 26 species from the families Chaetodontidae, Labridae and Pomacentridae were counted in 841 censuses; almost 80% of the fish belonged to just four species: bluehead (Thalassoma bifasciatum), bicolor damselfish (Stegastes partitus), slippery dick (Halichoeres bivittatus) and blue chromis (Chromis cyanea). Several multivariate techniques (cluster analysis, non‐metric multidimensional scaling and canonical correspondence analysis) were used to explore main patterns in assemblage composition variation. It was found that the results did not differ significantly when analyzing the same data set. Furthermore, it was shown that the combined use of different multivariate techniques enhanced the interpretation of fish assemblage composition changes. Depth was the main variable explaining variation in the composition of fish assemblages in the studied reefs. The densities of corals and gorgonians were also strongly associated with depth. Sponge density made an additional significant contribution to the explanatory model. Results of this research could be used as a baseline reference for future analyses of the impact of human actions in the study area. (e.g. oil spills, change in fishing intensity, other pollution events, tourism development).     

Species-level phylogeography and evolutionary history of the hyperdiverse marine gastropod genus Conus

Phylogenetic and paleontological analyses are combined to reveal patterns of species origination and divergence and to define the significance of potential and actual barriers to dispersal in Conus, a species-rich genus of predatory gastropods distributed throughout the world's tropical oceans. Species-level phylogenetic hypotheses are based on nucleotide sequences from the nuclear calmodulin and mitochondrial 16S rRNA genes of 138 Conus species from the Indo-Pacific, eastern Pacific, and Atlantic Ocean regions. Results indicate that extant species descend from two major lineages that diverged at least 33 mya. Their geographic distributions suggest that one clade originated in the Indo-Pacific and the other in the eastern Pacific + western Atlantic. Impediments to dispersal between the western Atlantic and Indian Oceans and the central and eastern Pacific Ocean may have promoted this early separation of Indo-Pacific and eastern Pacific + western Atlantic lineages of Conus. However, because both clades contain both Indo-Pacific and eastern Pacific + western Atlantic species, migrations must have occurred between these regions; at least four migration events took place between regions at different times. In at least three cases, incursions between regions appear to have crossed the East Pacific Barrier. The paleontological record illustrates that distinct sets of Conus species inhabited the Indo-Pacific, eastern Pacific + western Atlantic, and eastern Atlantic + former Tethys Realm in the Tertiary, as is the case today. The ranges of <1% of fossil species (N=841) spanned more than one of these regions throughout the evolutionary history of this group.     

Using Species-Area Relationships to Inform Baseline Conservation Targets for the Deep North East Atlantic

Demands on the resources of the deep-sea have increased in recent years. Consequently, the need to create and implement a comprehensive network of Marine Protected Areas (MPAs) to help manage and protect these resources has become a global political priority. Efforts are currently underway to implement MPA networks in the deep North East Atlantic. To ensure these networks are effective, it is essential that baseline information be available to inform the conservation planning process. Using empirical data, we calculated conservation targets for sessile benthic invertebrates in the deep North East Atlantic for consideration during the planning process. We assessed Species-Area Relationships across two depth bands (200–1100 m and 1100–1800 m) and nine substrata. Conservation targets were predicted for each substratum within each depth band using z-values obtained from fitting a power model to the Species-Area Relationships of observed and estimated species richness (Chao1). Results suggest an MPA network incorporating 10% of the North East Atlantic’s deep-sea area would protect approximately 58% and 49% of sessile benthic species for the depth bands 200–1100 m and 1100–1800 m, respectively. Species richness was shown to vary with substratum type indicating that, along with depth, substratum information needs to be incorporated into the conservation planning process to ensure the most effective MPA network is implemented in the deep North East Atlantic.