Ecological traits and environmental affinity explain Red Sea fish introduction into the Mediterranean

Alien species are considered one of the prime threats to biodiversity, driving major changes in ecosystem structure and function. Identifying the traits associated with alien introduction has been largely restricted to comparing indigenous and alien species or comparing alien species that differ in abundance or impact. However, a more complete understanding may emerge when the entire pool of potential alien species is used as a control, information that is rarely available. In the eastern Mediterranean, the marine environment is undergoing an unparalleled species composition transformation, as a flood of aliens have entered from the Red Sea following the opening of the Suez Canal in 1869. In this study, we compile data on species traits, geographical distribution, and environmental affinity of the entire pool of reef‐associated fish species in the Red Sea and more generally across the Indo‐Pacific. We use this extensive data to identify the prime characteristics separating Red Sea species that have become alien in the Mediterranean from those that have not. We find that alien species occupy a larger range of environments in their native ranges, explaining their ability to colonize the seasonal Mediterranean. Red Sea species that naturally experience high maximum temperatures in their native range have a high probability of becoming alien. Thus, contrary to predictions of an accelerating number of aliens following increased water temperatures, hotter summers in this region may prevent the establishment of many alien species. We further find that ecological trait diversity of alien species is substantially more evenly spaced and more divergent than random samples from the pool of Red Sea species, pointing at additional processes, such as competition, promoting ecological diversity among alien species. We use these results to provide a first quantitative ranking of the potential of Red Sea species to become established in the eastern Mediterranean.     

Population status of giant clams (Mollusca: Tridacnidae) in the northern Red Sea,Egypt

Throughout their range, giant clams (family Tridacnidae) are increasingly threatened by anthropogenic impacts and natural disasters, but little is known about their population status. In this first assessment of the tridacnid population at Abu Sauatir in the northern Red Sea, a total of 491 m² were surveyed and >200 clams recorded. Tridacna maxima was the only species found. The population's live:dead ratio was 3:1. Overall clam density was 0.08?±?0.008 live and 0.02?±?0.007 dead individuals per 0.25 m². Greatest densities occurred on the reef flat in 1 m depth (live), and on the northern reef slope in deeper waters (dead). On the slope, live clam density decreased significantly, whereas dead clam density increased significantly with depth. Sizes of live and dead individuals differed significantly. Live clams ranged from 1 to 30 cm (median 5 cm). Juveniles ≤2 cm (8.2% of the population) and individuals >11 cm occurred on the reef slope but not on the reef flat. Live clam sizes did not differ significantly between reef sites. Dead clam size ranged from 2 to 15 cm (median 6.5 cm). 2.1% of the empty shells were ≤2 cm long. Dead clam sizes differed significantly between 5 and 10 m depth on the northern reef slope. The low clam abundance (live and dead) in the shallowest and most easily accessible areas of the reef flat, combined with small sizes, strongly suggest artisanal reef-top gathering for meat and shells.     

Bidirectional sex-change behavior and physiological aspects in the Gorgeous goby Lythrypnus pulchellus (Gobiidae)

The Gorgeous goby Lythrypnus pulchellus shows extreme sexual plasticity with the bidirectional sex-change ability socially controlled in adults. Therefore, this study describes how the hierarchical status affects hormone synthesis through newborn hormone waste products in water and tests the influence of body size and social dominance establishment in sex reversal duration and direction. The associated changes in behavior and hormone levels are described under laboratory conditions in male–male and female–female pairs of similar and different body sizes, recording the changes until spawning. The status establishment occurred in a relatively shorter time period in male and female pairs of different sizes (1–3 days) compared to those of similar size (3–5 days), but the earlier one did not significantly affect the overall time of sex change (verified by pair spawning). The changes in gonads, hormones, and papilla occurred in sex-changer individuals, but the first one was observed in behavior. Courtship started at 3–5 days in male pairs and from 2 h to 1 day in female pairs of both groups of different and similar sizes. Hormones did not gradually move in the new sexual phenotype direction during the sex-change time course. Nonetheless, estradiol regulated sex change and 11-ketotestosterone enabled bidirectional sex change and was modulated by agonistic interactions. Cortisol is associated with status and gonadal sex change. In general, similar mechanisms underlie sex change in both directions with a temporal change sequence in phases. These results shed new light on sex-change mechanisms. Further studies should be performed to determine whether these localized changes exist in the steroid hormone synthesis along the brain–pituitary gonad axis during social and bidirectional sex changes in L. pulchellus.     

Comparison of deep-water coral reefs and lithoherms off southeastern USA

Two types of deep-water coral bioherms occur off the coast of southeastern United States: Oculina and Lophelia/Enallopsammia. The deep-water Oculina bioherms form an extensive reef system at depths of 70–100 m along the shelf edge off central eastern Florida. These reefs are comprised of numerous pinnacles and ridges, 3–35 m in height. Each pinnacle is a bank of unconsolidated sediment and coral debris that is capped on the slopes and crest with living and dead colonies of Oculina varicosa, the ivory tree coral. In comparison, deep-water reefs of Lophelia pertusa and Enallopsammia profunda corals occur at depths of 500–850 m (maximum 150-m relief) along the base of the Florida-Hatteras slope in the Straits of Florida. On the western edge of the Blake Plateau off South Carolina and Georgia, 54-m high banks of Enallopsammia and Lophelia occur at depths of 490–550 m, whereas on the eastern edge of the plateau the reefs form structures 146 m in height and at depths of 640–869 m. The geomorphology and functional structure of both the Oculina and Lophelia reefs are similar. North of Little Bahama Bank, at depths of 1000–1300 m, a region of bioherms is dominated by the coral Solenosmilia sp.; Lophelia is reportedly absent. This paper summarizes 25 years of submersible studies on the deep-water Oculina reefs, describes submersible reconnaissance of deep-water Lophelia reefs off the southeastern United States, and contrasts these types of bioherms with the deep-water lithoherms in the Straits of Florida west of the Bahamas.     

Dating the evolutionary origins of wrasse lineages (Labridae) and the rise of trophic novelty on coral reefs

We estimated ages of divergence between major labrid tribes and the timing of the evolution of trophic novelty. Sequence data for 101 labrid taxa and 14 outgroups consisting of two mitochondrial gene regions (12s, 16s), and two nuclear protein-coding genes (RAG2, TMO4c4), a combined 2567 bp of sequence, were examined using novel maximum likelihood, maximum parsimony and mixed model Bayesian inference methods. These analyses yielded well supported trees consistent with published phylogenies. Bayesian inference using five fossil calibration points estimated the minimum ages of lineages. With origins in the late Cretaceous to early tertiary, the family diversified quickly with both major lineages (hypsigenyine and julidine) present at approximately 62.7 Ma, shortly after the K/T boundary. All lineages leading to major tribes were in place by the beginning of the Miocene (23 Ma) with most diversification in extant lineages occurring within the Miocene. Optimisation of trophic information onto the chronogram revealed multiple origins of novel feeding modes with two distinct periods of innovation. The Palaeocene/Eocene saw the origins of feeding modes that are well represented in other families: gastropod feeders, piscivores and browsing herbivores. A wave of innovation in the Oligocene/Miocene resulted in specialized feeding modes, rarely seen in other groups: coral feeding, foraminifera feeding and fish cleaning. There is little evidence of a general relationship between trophic specialization and species diversity. The current trophic diversity of the Labridae is a result of the accumulation of feeding modes dating back to the K/T boundary at 65 Ma, with all major feeding modes on present day reefs already in place 7.5 million years ago.     

Thermoregulatory behavior of the triggerfish Balistes fuscus in an electronic shuttlebox

Ten blue triggerfish,Balistes fuscus, were tested individually for 3 days each in Ichthyotron electronic shuttleboxes to measure their thermoregulatory behavior. The modal thermal preferendum, a species-specific measure of temperature preference which is independent of prior thermal acclimation, was 25 °C. The triggerfish voluntarily occupied a 16–27 °C range of temperature, out of a potentially available range of 0–50 °C. There was no significant difference in preferred temperature between night and day, indicating lack of a thermoregulatory rhythm in this species. The preferred temperature range of this tropical marine reef species is similar to that of cool temperate freshwater and marine fishes; many warm temperate species prefer higher temperatures.     

Nocturnal relocation of adult and juvenile coral reef fishes in response to reef noise

Juvenile and adult reef fishes often undergo migration, ontogenic habitat shifts, and nocturnal foraging movements. The orientation cues used for these behaviours are largely unknown. In this study, the use of sound as an orientation cue guiding the nocturnal movements of adult and juvenile reef fishes at Lizard Island, Great Barrier Reef was examined. The first experiment compared the movements of fishes to small patch reefs where reef noise was broadcast, with those to silent reefs. No significant responses were found in the 79 adults that were collected, but the 166 juveniles collected showed an increased diversity each morning on the reefs with broadcast noise, and significantly greater numbers of juveniles from three taxa (Apogonidae, Gobiidae and Pinguipedidae) were collected from reefs with broadcast noise. The second experiment compared the movement of adult and juvenile fishes to reefs broadcasting high (>570 Hz), or low (<570 Hz) frequency reef noise, or to silent reefs. Of the 122 adults collected, the highest diversity was seen at the low frequency reefs; and adults from two families (Gobiidae and Blenniidae) preferred these reefs. A similar trend was observed in the 372 juveniles collected, with higher diversity at the reefs with low frequency noises. This preference was seen in the juvenile apogonids; however, juvenile gobiids were attracted to both high and low sound treatments equally, and juvenile stage Acanthuridae preferred the high frequency noises. This evidence that juvenile and adult reef fishes orientate with respect to the soundscape raises important issues for management, conservation and the protection of sound cues used in natural behaviour.     

Effects of habitat-conditioning by the damselfish Stegastes nigricans (Lacepède) on the community structure of benthic algae

We compared the community structure of benthic algae inside and outside pomacentrid damselfish (Stegastes nigricans) territories in a moat at Sesoko Island, Okinawa, Japan. S. nigricans maintained “algae farms” that were dominated by the filamentous rhodophyte, Womersleyella setacea. Species richness and biomass were higher inside damselfish territories than outside, while species diversity and evenness were higher outside. Detrended correspondence analysis (DCA) based on species composition showed that the dominance of W. setacea was maintained throughout the year in all samples collected from inside damselfish territories. The observed strong dominance of filamentous rhodophytes was consistent with the findings of most studies on damselfish territories worldwide. However, the dominance of a single species of alga and low species diversity inside the territories was in contrast to the findings of previous studies, in which the reduction of grazing pressure caused intermediate disturbance and enhanced algal species diversity. This discrepancy in algal species diversity inside the damselfish territories seems to have been caused by unique characteristics of the alga and the fish. W. setacea traps sediment, which reduces the availability of firm substrata for attachment and inhibits the recruitment of some algae. Moreover, S. nigricans “weeds” indigestible calcareous and thicker algae. The algal assemblage outside damselfish territories varied among samples, and included mat-forming cyanophytes (Calothrix aeruginosa and Calothrix codicola), a prostrate laminar phaeophyte (Padina sp.), thin and small-scaled algae (Cladophora sp. and Feldmannia indica), finely branched filamentous rhodophytes (Taenioma perpusillum and Herposiphonia obscura), and a coarsely branched rhodophyte (Gelidiopsis variabilis).

We placed artificial slate plates inside and outside damselfish territories, and showed that the W. setacea inside territories gradually increased in biomass, reaching the same levels of biomass and dominance as W. setacea on natural substrata. Outside the territories, the algal assemblage underwent succession from early colonizers, i.e., thin and small-scaled algae, to grazing-resistant algae such as mat-forming cyanophytes and prostrate laminar Padina sp. Under heavy grazing, the flora outside the territories was composed of early colonizers, grazing-resistant algae, and scattered erect algae that had probably escaped grazing by chance.

Our findings suggest that sediment trapped by the turf of W. setacea inhibited recruitment of some algae, and that moderate cropping and selective weeding by S. nigricans excluded grazing-resistant algae and prevented early colonizers and competitively superior algae from out-competing W. setacea. Consequently, low species diversity and a high-biomass “farm” suitable for harvesting was maintained.     

Effects of low light and high temperature on pediveligers of the fluted giant clam Tridacna squamosa

ABSTRACT

This paper examines the responses of the fluted giant clam Tridacna squamosa pediveligers to elevated temperature and reduced light levels. In a light reduction experiment, a total of 104,000 T. squamosa pediveligers were exposed to four different levels of shading for approximately one month. The most heavily shaded treatment, at 0.4% of ambient light, had significantly lower survival than the other groups, which all received 1% or more of ambient light. In a second experiment, for approximately two weeks 13,000 T. squamosa pediveligers were divided among three treatments: one at ambient temperature averaging 29.5 °C, and two with elevated temperatures averaging 32.2 °C and 34.8 °C. The elevated temperature treatments resulted in near total mortality. The highest temperature survived by any pediveliger was 32.8 °C. Our results indicate a potential synergetic effect, with turbidity causing giant clam pediveligers to settle in shallower water―where they will likely be exposed to higher temperatures.     

A new sponge-inhabiting amphipod species (Crustacea, Gammaridea, Sebidae) from the Veracruz Coral Reef System, southwestern Gulf of Mexico

A new species of the genus Seba Bate (Amphipoda: Sebidae) is described from the Veracruz Coral Reef System, Veracruz, south-western Gulf of Mexico. The specimens were found in association with a sponge, Ircinia fistularis, at depths from 8 to 10 m. Seba alvarezi n. sp. can be distinguished from closely related species mainly by the presence of a short, two-articulated accessory flagellum, three apical setae on article 3 of the mandible palp, and by the absence of apical setae on the inner lobe of maxilla 1. The new species is compared to closely related species in the family, and an identification key to the species of Seba known from the Gulf of Mexico and the Caribbean Sea is given.