Trophic specialization influences the rate of environmental niche evolution in damselfishes (Pomacentridae)

The rate of environmental niche evolution describes the capability of species to explore the available environmental space and is known to vary among species owing to lineage-specific factors. Trophic specialization is a main force driving species evolution and is responsible for classical examples of adaptive radiations in fishes. We investigate the effect of trophic specialization on the rate of environmental niche evolution in the damselfish, Pomacentridae, which is an important family of tropical reef fishes. First, phylogenetic niche conservatism is not detected in the family using a standard test of phylogenetic signal, and we demonstrate that the environmental niches of damselfishes that differ in trophic specialization are not equivalent while they still overlap at their mean values. Second, we estimate the relative rates of niche evolution on the phylogenetic tree and show the heterogeneity among rates of environmental niche evolution of the three trophic groups. We suggest that behavioural characteristics related to trophic specialization can constrain the evolution of the environmental niche and lead to conserved niches in specialist lineages. Our results show the extent of influence of several traits on the evolution of the environmental niche and shed new light on the evolution of damselfishes, which is a key lineage in current efforts to conserve biodiversity in coral reefs.     

Isolation and characterization of 16 microsatellite loci in the humbug damselfish, Dascyllus aruanus (family Pomacentridae)

Here we report on 16 microsatellite loci designed for the damselfish Dascyllus aruanus. All loci were tested on 98 individuals and were polymorphic (seven to 35 alleles). Expected heterozygosity ranged from 0.705 to 0.942. Six loci showed Hardy-Weinberg disequilibrium due to the occurrence of null alleles. Cross-species amplifications conducted within the genus Dascyllus (D. carneus, D. strasburgi, D. trimaculatus) lead to polymorphic fragments in 32 out of 48 tests. These 16 loci will enable future research into the behavioural ecology and population ecology of Dascyllus aruanus throughout the Indo-Pacific.     

Trait decoupling promotes evolutionary diversification of the trophic and acoustic system of damselfishes

Trait decoupling, wherein evolutionary release of constraints permits specialization of formerly integrated structures, represents a major conceptual framework for interpreting patterns of organismal diversity. However, few empirical tests of this hypothesis exist. A central prediction, that the tempo of morphological evolution and ecological diversification should increase following decoupling events, remains inadequately tested. In damselfishes (Pomacentridae), a ceratomandibular ligament links the hyoid bar and lower jaws, coupling two main morphofunctional units directly involved in both feeding and sound production. Here, we test the decoupling hypothesis by examining the evolutionary consequences of the loss of the ceratomandibular ligament in multiple damselfish lineages. As predicted, we find that rates of morphological evolution of trophic structures increased following the loss of the ligament. However, this increase in evolutionary rate is not associated with an increase in trophic breadth, but rather with morphofunctional specialization for the capture of zooplanktonic prey. Lineages lacking the ceratomandibular ligament also shows different acoustic signals (i.e. higher variation of pulse periods) from others, resulting in an increase of the acoustic diversity across the family. Our results support the idea that trait decoupling can increase morphological and behavioural diversity through increased specialization rather than the generation of novel ecotypes.     

Evolutionary history of the butterflyfishes (f: Chaetodontidae) and the rise of coral feeding fishes

Of the 5000 fish species on coral reefs, corals dominate the diet of just 41 species. Most (61%) belong to a single family, the butterflyfishes (Chaetodontidae). We examine the evolutionary origins of chaetodontid corallivory using a new molecular phylogeny incorporating all 11 genera. A 1759‐bp sequence of nuclear (S7I1 and ETS2) and mitochondrial (cytochrome b) data yielded a fully resolved tree with strong support for all major nodes. A chronogram, constructed using Bayesian inference with multiple parametric priors, and recent ecological data reveal that corallivory has arisen at least five times over a period of 12 Ma, from 15.7 to 3 Ma. A move onto coral reefs in the Miocene foreshadowed rapid cladogenesis within Chaetodon and the origins of corallivory, coinciding with a global reorganization of coral reefs and the expansion of fast‐growing corals. This historical association underpins the sensitivity of specific butterflyfish clades to global coral decline.     

Four new species of dwarfgobies (Teleostei: Gobiidae: Eviota) from the Austral,Gambier, Marquesas and Society Archipelagos,French Polynesia

Four new species of Eviota (Teleostei: Gobiidae) are described from French Polynesia. Eviota hinanoae occurs from the Gambier, Austral and Society Archipelagos and differs from its closest relatives primarily in sensory pore pattern, having cuplike male urogenital papilla, and in lacking prominent dark pigmentation on the base of the pectoral fin. Three other new species, E. dorsimaculata, E. lacrimosa and E. deminuta, are known only from the Marquesas Islands and are distinguished from congeners by differences in pigmentation, sensory pore pattern, urogenital papilla shape, the number of rays in the dorsal and anal fins, and the length of the fifth pelvic-fin ray. Eviota deminuta represents one of the smallest known species of Eviota and features reductions in several morphological characters, including the loss of all head pores. Three of the new species are included in a phylogenetic analysis based on mitochondrial cytochrome c oxidase-I, along with 21 additional species of Eviota.     

Molecular phylogeny of the entomopathogenic fungi of the genus Cordyceps (Ascomycota:Clavicipitaceae) and its evolutionary implications

Cordyceps is an endoparasite ascomycetous genus containing approximately 450 species with a diversity of insect hosts,traditionally included in the family Clavicipitaceae of Ascomycota.Establishing the relationships among species with a varied range of morphologies and hosts is of importance to our understanding of the phylogeny and co-evolution of parasites and hosts in entomopathogenic ascomycetes.To this end,we used a combination of molecular index and morphological characters from 40 representative species to carry out comprehensive molecular phylogenetic analyses.Based on the phylogenetic tree,we used the program DISCRETE for inferring the rates of evolution and finding ancestral states of morphological character.The phylogenetic analyses revealed two important points.(i) Types of perithecia attached to stroma reflected an evolutionary trend in Cordyceps.The vertically immersed perithecia form was the ancestral state,superficial and obliquely immersed perithecia were derived characters,obliquely immersed was irreversible.Species with obliquely immersed perithecia were in a closely related group and were the derived group.(ii) A strong correlation between fungal relatedness and the microhabitat supported the hypothesis that the host jumps through commingling in soil microhabitats.Based on the results of these analyses,host switching explains the diversity of entomopathogenic fungi of the genus Cordyceps.     

Correlated evolution of sex and reproductive mode in corals (Anthozoa: Scleractinia)

Sexuality and reproductive mode are two fundamental life-history traits that exhibit largely unexplained macroevolutionary patterns among the major groups of multicellular organisms. For example, the cnidarian class Anthozoa (corals and anemones) is mainly comprised of gonochoric (separate sex) brooders or spawners, while one order, Scleractinia (skeleton-forming corals), appears to be mostly hermaphroditic spawners. Here, using the most complete phylogeny of scleractinians, we reconstruct how evolutionary transitions between sexual systems (gonochorism versus hermaphrodism) and reproductive modes (brooding versus spawning) have generated large-scale taxonomic patterns in these characters. Hermaphrodites have independently evolved in three large, distantly related lineages consisting of mostly reef-building species. Reproductive mode in corals has evolved at twice the rate of sexuality, while the evolution of sexuality has been heavily biased: gonochorism is over 100 times more likely to be lost than gained, and can only be acquired by brooders. This circuitous evolutionary pathway accounts for the prevalence of hermaphroditic spawners among reef-forming scleractinians, despite their ancient gonochoric heritage.     

Small-scale spatial variation in the elemental composition of otoliths of Stegastes nigricans (Pomacentridae) in French Polynesia

Solution-based inductively coupled plasma-mass spectrometry (ICP-MS) was used to determine if Stegastes nigricans collected from 15 sites in French Polynesia could be distinguished by the trace element composition of their otoliths. A total of 293 adults were collected by spearing and their otoliths were analysed. We found that elemental signatures differed significantly among sites within and between the islands of Tahiti and Moorea (p<0.001), primarily due to variation in concentrations of the elements Ba, Ca, Li, Mg, Mn, Na, Sr and Y. The otoliths of fish collected within Papeete Harbour in Tahiti had distinctive elemental signatures characterised by relatively high concentrations of Mn. Otoliths of these fish could be distinguished from others that were collected only a small distance (200 m) from the harbour. This is the first time that differences in chemical composition of otoliths have been reported at such small spatial scales and this trait may prove useful for the studies of connectivity of populations at within reef scales.     

Colonial Nesting and the Importance of the Brood Size in Male Parasitic Reproduction of the Mediterranean Damselfish Chromis chromis (Pisces: Pomacentridae)

We investigated male parasitic spawning in a protected natural population of Mediterranean damselfish. Chromis chromis nested in colonies, inside which males showed a high variability in mating success. Our field observations indicate that the egg batches obtained by the most successful fish were five times bigger than the ones obtained by the less successful fish and many males never received ovipositions. On the other hand, reproductive parasitism was a common tactic within the colony. Successful nesting males sneaked into their neighbours' nests depending on the amount of eggs in their nest, with small clutch size inducing the males to parasitic reproduction. Males failing to receive egg depositions on their nests showed a significantly higher parasitism rate than successful males. Non-territorial males occupied stations in the water column above the breeding grounds and whenever the opportunity arose, they disrupted spawning in progress, stealing copulation with females. We observed that the likelihood of males being parasitized by sneakers was not correlated with the size of their own clutch; on the contrary, it depended both on the number of neighbouring nests and on the number of neighbouring males with barren nests (i.e. unsuccessful males). No correlation was found between parasitic behaviour and male size, suggesting males may switch between spawning in their own and in their neighbour's nests depending on mating opportunity. The hypothesis that colonial nesting facilitates parasitic reproduction is here discussed.     

Geological history and oceanography of the Indo‐Malay Archipelago shape the genetic population structure in the false clown anemonefish (Amphiprion ocellaris)

Like many fishes on coral reefs, the false clown anemonefish, Amphiprion ocellaris, has a life history with two different phases: adults are strongly site attached, whereas larvae are planktonic. Therefore, the larvae have the potential to disperse, but the degree of dispersal potential depends primarily on the period of the larval stage, which is only 8–12 days in A. ocellaris. In this study, we investigated the genetic population structure and gene flow in A. ocellaris across the Indo‐Malay Archipelago by analysing a fragment of the mitochondrial control region. Population genetic analysis, using amova , revealed a significant and high overall Φ_ST‐value of 0.241 (P < 0.001), clearly showing limited gene flow. Haplotype network analysis detected eight distinct clades corresponding mainly to different geographical areas, which were most probably separated during sea level low stands in the Pleistocene. The distribution of the clades among the different populations indicated slow partial re‐mixing mainly in the central region of the archipelago. Major surface currents seem to facilitate larval dispersal, indicated by higher connectivity along major surface currents in the region (e.g. Indonesian Throughflow). Four main groups were found by the hierarchical amova within the archipelago. These different genetic lineages should be managed and protected as separate ornamental fishery stocks and resource contributing to the genetic diversity of the area. Regarding the high diversity and the differentiation among areas within the Indo‐Malay Archipelago of A. ocellaris populations, the centre‐of‐origin theory is supported to be the main mechanism by which the high biodiversity evolved in this area.     

Interspecific variation in sustained swimming ability of late pelagic stage reef fish from two families (Pomacentridae and Chaetodontidae)

 Characteristics of the pelagic stages of reef fishes have generally been investigated at the family level, which may mask important differences among species. Here the variation in sustained swimming ability of the late pelagic stages is examined among species, within two families (Pomacentridae and Chaetodontidae). The pomacentrids displayed a 7.5-fold difference in sustained swimming ability across 24 species, while the chaetodontids displayed a 2-fold difference across 10 species. The variation within the Pomacentridae was not related to pelagic larval duration, post-settlement habitat or taxonomy. There was, however, a significant correlation between sustained swimming ability and total length (TL) of individuals (r=0.435, P<0.0001). Differences in the mean distance swum by pomacentrid species, however, was most strongly related to differences in mean wet weight (r=0.814, P<0.0001). When the mean distance swum by species was scaled with respect to mean TL there was still a strong correlation with mean wet weight (r=0.644, P<0.005). Among chaetodontid individuals TL and sustained swimming ability were not correlated (r=−0.004, P=0.978). Furthermore, sustained swimming ability was not significantly related to the trans-oceanic distribution of species in either family. The variation in sustained swimming ability, however, may contribute to explanations of the observed levels of gene flow within populations. Accepted: 24 November 1997     

A review of the fossil record of the Pomacentridae (Teleostei: Labroidei) with a description of a new genus and species from the Eocene of Monte Bolca, Italy

The fossil record of the Pomacentridae has been based on species from six genera. The status of these taxa is reviewed. Only two are tentatively accepted as belonging to the Pomacentridae: Chromis savomini from the Miocene of Algeria and Izuus nakamurai from the Miocene of Japan. In addition, a new genus and species Palaeopomacentrus orphae is described from the lower Middle Eocene of Monte Bolca, Northern Italy. This is the first substantiated record of a pomacentrid from the Eocene, and represents the oldest record of the family.     

Correlates of Environmental Variables with Patterns in the Distribution and Abundance of Two Anemonefishes (Pomacentridae: Amphiprion) on an Eastern Australian Sub-tropical Reef System

The relationship between characteristics of the reef environment and variations in the distribution and abundance of the anemonefishes Amphiprion akindynos and A. latezonatus was investigated at North Solitary Island, a sub-tropical rocky reef system on the east coast of Australia. During the summers of 1994 and 1995, fish densities and host sea-anemone cover were assessed on replicate 25m transects at sites where host sea-anemones form semi-contiguous mats throughout the 6–21m depth range. Multiple regression analyses indicated that environmental variables accounted for 65–71% and 61–80% of the variations in the number of A. akindynos and A. latezonatus among-sites, respectively. Among-habitat comparisons indicated that A. latezonatus densities were positively correlated with depth (r=0.45–0.90), whereas A. akindynos showed no consistent depth-related abundance patterns. Poor correlations (p>0.05) between the densities of each species on transect lines suggested that present-day competition was unlikely to determine the preference of A. latezonatus for deeper depths. Correlations between host sea-anemone cover and fish densities at the within-habitat (depth) scale were comparatively stronger than correlations at among-habitats in both species. These results suggest that among-habitat comparisons can confound finer scale fish-habitat associations within habitat (depth) zones. Evidence suggests that while sea-anemone cover does, to an extent, regulate the local ecology of anemonefishes, other factors are also likely to interact to limit their densities.     

Spatio-temporal patterns of recruitment of labroid fishes (Pisces: Labridae and Scaridae) to damselfish territories

Recent studies have shown that recruitment variability is an important process structuring reef fish assemblages. The aim of this study is to describe the spatio-temporal patterns of recruitment of three abundant labroid taxa (Coris schroederi, Halichoeres melanurus and Scarus spp.), using damselfish territories as replicate units of habitat. Temporal recruitment patterns of each taxa were consistent among three sites along 2 km of reef tract, with small differences among sites probably the result of hydrological or random factors operating at that scale. Recruitment of only one species (C. schroederi) showed consistent differences in the magnitude of recruitment among sites, which was probably due to an overriding effect of habitat selection for the location of the territories on the reef profile at one site. Two taxa, C. schroederi and Scarus spp., recruited in low to moderate rates over many weeks with moderate recruitment peaks detected in one year only. This pattern may be characteristic of many labroid species that have protracted periods of production of larvae. In contrast, H. melanurus recruited in a single short pulse of high magnitude each summer, which suggests that production of larvae by this species may occur over a more restricted period of time. No strong pattern of lunar entrainment of recruitment was detected for any taxa, which may be due to a lack of lunar periodicity in production of larvae. Further studies are now required to identify the processes that are important in determining patterns of labroid recruitment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Aggregations of <Emphasis Type="Italic">Plectropomus areolatus</Emphasis> and <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (groupers,Serranidae) in the Komodo National Park,Indonesia: Monitoring and Implications for Management

Synopsis We identify fishery management implications from a long-term monitoring program focusing on spawning aggregations of high valued reef fish in Komodo National Park (KNP), Eastern Indonesia. Management objectives of KNP are not only to protect biodiversity, but also to conserve spawning stocks of high-valued commercial species for the replenishment of surrounding fishing grounds. We monitored two sites twice monthly over five years for two species of grouper, Epinephelus fuscoguttatus and Plectropomus areolatus. One site had an aggregation of both E. fuscoguttatus and P. areolatus, whereas the other site contained an aggregation of P. areolatus only. Over the five years monitoring period, aggregations typically formed during each full moon between September and February. Additionally, P. areolatus occasionally aggregated during new moons between April and July. We observed spawning only once, but because formation of aggregations were correlated to a higher incidence of behavior and signs indicative of reproduction and because most fish present were adults, it is likely that the formation of aggregations was associated with spawning. Over the five years monitoring period there was a reduction in mean fish size of up to 8 cm for P. areolatus, and a reduction in numbers of aggregating E. fuscoguttatus. Despite limited protection initiated in 2001, both sites are still heavily fished by local artisanal fishers. Because the observed reductions in size and in numbers could be caused by fishing pressure, managers should follow the precautionary principle by putting additional protective management in place. Since both species are relatively long-lived, at least five years of continued monitoring may be necessary to determine the outcome of management intervention. The variability in timing of aggregation in respect to season and moon phase in P. areolatus indicates that long-term monitoring must cover the entire year and both moon phases.     

Malaria parasites (Apicomplexa,Haematozoea) and their relationships with their hosts: is there an evolutionary cost for the specialization?

Parasite–host specialization is frequently considered to be a derived state such that it represents an 'evolutionary dead end' that strongly limits further evolution. In this study, it was tested whether this theory is applicable to the relationship of malaria parasites and their vertebrate hosts. For this, we revisited Perkins and Schall (2002) analysis of the phylogenetic relationships of the malaria parasites (belonging to the genera Plasmodium , Haemoproteus and Hepatocystis ) based on the mitochondrial Cytochrome b gene sequence, and inferred, using a maximum likelihood (ML) approach, the putative ancestral vertebrate hosts. As the topology in this study presents several unresolved branches and is slightly different from that of Perkins and Schall, a Shimodaira and Hasegawa (SH; 1999) test has been performed in order to properly consider several alternative topologies. The results of this study suggest that the common ancestor of all these malaria parasites was a reptile (more specific of the order Squamata), and that the host switches from Squamata to Aves and vice versa were quite frequent along the evolution of these parasites. On the contrary, a strong evidence that the host shift from Squamata to Mammalia had occurred only once during the evolution of these organisms was found. This evidence (added to the current knowledge about the association of the malaria parasites with their vertebrate hosts) allows us to suggest, at least considering the species included in this study, that the adaptation in mammals had required a high level of specialization. Hence, the acquisition of this host class had culminated in an evolutionary dead end for the mammalian malaria parasites.     

Systenostrema alba Larsson 1988 (Microsporidia, Thelohaniidae) in the Dragonfly Aeshna viridis (Odonata, Aeshnidae) from South Siberia: morphology and molecular characterization

An octospore microsporidium was found in the nymphs of Aeshna viridis, collected in intermittent streams near Novosibirsk, Siberia, Russia in 2003. Spores were uninucleate and measured 6.1+/-0.07 x 3.0+/-0.04 microm on fresh smears. The polar filament was anisofilar having 10-11 anterior coils (thicker filament diam.) and 10-11 posterior (thinner filament diam.) coils. Sporophorous vesicles were persistent and measured 12.3+/-0.23 x 11.9+/-0.20 microm. The infection was restricted to the adipose tissue and caused the formation of whitish "cysts" containing mature octospores. Based on ultrastructural similarity we consider this Siberian isolate to be Systenostrema alba, a species described from Aeshna grandis collected in Sweden (Larsson 1988). Maximum likelihood, neighbor joining, and maximum parsimony analyses of the small subunit rDNA all placed Systenostrema alba (Accession no. AY953292) as the sister taxon to a clade consisting of Thelohania solenopsae, Tubulinosema ratisbonensis, and Tubulinosema acridophagus.     

Some Evolutionary Arguments about what maintains the Pelagic Interval in Reef Fishes

In this paper, we address the question of the nature of evolutionary forces that account for the oceanic planktonic larva period. Putting the emphasis on coral reef fishes, we first present the most common theories and hypotheses, looking at them critically from the standpoint of individual selection and taking into account new data arising from genetic surveys. We concluded that each individual hypothesis based on short term advantages of a larva period cannot convincingly explain by itself the long term maintenance of such a complex life cycle. We then study the impact of the pelagic interval on species dispersal by compiling data sets from the literature. Following simple analysis of this data, we found that duration of the planktonic larva period drives gene flow in the Great Barrier Reef and colonisation throughout the Pacific. In speculating on the real nature of the short term selective forces responsible for the maintenance of the pelagic interval, we acknowledge the fact that long term constraints alone will not withstand erosion if they are not reinforced by some sort of short term mechanism. We tentatively arrive at the conclusion that these short term forces may be different from what could be expected from functional studies of planktonic life.     

Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida)

The phylogenetic relationships of the Clitellata were investigated using a data set with published and new complete or partial 18S rRNA and mtCOI gene sequences of 13 and 49 taxa representing 8 and 14 families, respectively. Three different alignments were considered for 18S, and the possible influence of departures from rate constancy among sites was evaluated by analyses using a Gamma model of rate heterogeneity. Maximum-likelihood estimates of the shape parameter alpha of the Gamma distribution were very low, whatever the alignment or the gene considered, suggesting that phylogenetic reconstructions taking into account the rate heterogeneity among sites are likely to be the most reliable. Analyzed separately, the two genes did not resolve the relationships among the Clitellata, but the consensus tree was congruent with the morphology-based relationships. Our data suggest the inclusion of the Euhirudinea, Acanthobdellida, and Branchiobdellida in the Oligochaeta and suggest the Lumbriculidae as the link between both assemblages. Although separate analyses of both genes, as well as different alignments for the 18S rRNA sequences, yielded conflicting results concerning the phylogenetic position of leeches and leech-like worms vis-à-vis the Oligochaeta, subsequent analyses using the Gamma model greatly reduced the observed inconsistencies. Our analyses show that among the Clitellata, the leeches and the leech-like and gutless worms represent significantly faster evolving lineages. It is suggested that the observed higher mutation rates may be explained by the fact that these lineages contain almost exclusively commensal and/or parasitic taxa.