The damselfishes (Pisces: Osteichthyes: Pomacentridae) of Peninsular Malaysia and Singapore: systematics,ecology and conservation

Damselfishes (Pomacentridae) are among the most common of reef fishes in tropical seas, comprising 320 species in all, with 268 species alone in the Indo-West Pacific regions. 15 genera (Abudefduf, Amblyglyphidodon, Amphiprion, Chelioprion, Chromis, Chrysiptera, Dascyllus, Pristotis, Dischistodus, Hemiglyphidodon, Neoglyphidodon, Neopomacentrus, Plectroglyphidodon, Pomacentrus, Stegastes) and at least 39 species inhabit the waters of Peninsular Malaysia and Singapore. They are important ecologically because many species are extremely abundant in reefs, and also economically, as several are collected in large numbers for the lucrative aquarium trade. This demand has led to some fishermen using destructive methods in obtaining the fishes, to the point that original standing populations may not be recoverable. This threat poses a need for conservation of the reefs. The present study based primarily on fishes collected from the waters of Singapore and specimens from the Zoological Reference Collection at the National University of Singapore, will serve the dual purpose of establishing a current species list for Malaysia and Singapore, and to update and clarify the local taxonomy of the group. An annotated list (with localities) is provided of the known or reported species to aid studies on reefs.     

Biosociology and ecology of pomacentrid fishes around the Sinai Peninsula (northern Red Sea)

Pomacentrid fishes inhabiting the coral formations along the Red Sea coast of Sinai Peninsula form specific groups of species on various habitats. Amphirion bicinctus, Dascyllus aruanus, D. marginatus and Pomacentrus trichourus are typical for the backreef lagoon; Pomacentrus sulfureus, Abudefduf melanopus and A. melas are common along the rearreef wall; Pomacentrus albicaudatus, P. tripunctatus, Abudefduf lacrymatus, A. leucozona and A. annulatus dominate the coral reef table. The forereef wall is occupied by Abudefduf saxatilis, A. sexfasciatus and A. leucogaster . The lower part of the forereef and coral knolls along it are inhabited by groups of Abudefduf azysron, Chromis dimidiatus, C. caeruleus, C. ternatensis and of the Dascyllus trimaculatus . Among the fishes observed a part are living in groups or schools, whereas others are typical solitary, permanent territory holders. Most of them are polygamous, breeding in colonies, or solitary in nests, guarded by males. The variation in feeding behaviour as well as in sociobiology, enables the high diversity of these fishes on various parts of the habitat.     

Predation on coral spawn by planktivorous fish

The stomach contents of three species of coral reef fish were examined for changes in diet during the annual mass spawning of scleractinian corals on the Great Barrier Reef, Australia. Acanthochromis polyacanthus, Abudefduf bengalensis (Pomacentridae), and Caesio cuning (Caesionidae) were collected before and immediately after the coral spawning to determine whether the composition of the diet changed after the mass coral spawning. The diet of Caesio cuning did not change. The stomach contents of Acanthochromis and Abudefduf showed that these fish: (1) switched from an omnivorous diet to one consisting predominantly of coral spawn, and (2) that they appeared satiated with stomachs >90% full when coral spawn was present in the water column. Fish predation during coral spawning may be an important source of larval coral mortality.     

三亚珊瑚礁保护区珊瑚礁生态系统现状及其健康状况评价

为评估三亚珊瑚礁国家级自然保护区珊瑚礁生态系统的健康状况, 本文选取东岛、鹿回头、大东海3个站位调查了珊瑚礁群落、珊瑚礁鱼类和大型底栖动物。通过对比分析历史资料、珊瑚礁现场生态调查与监测及组织专家评审, 筛选出一、二级指标并设置权重, 使用综合指数计算了三亚珊瑚礁保护区珊瑚礁生态系统健康指数。结果显示, 三亚珊瑚礁保护区内共有造礁珊瑚10科21属37种, 软珊瑚3种, 造礁珊瑚覆盖率和软珊瑚覆盖率分别为14.31%和0.19%, 其中鹿回头造礁珊瑚覆盖率最高, 为21.58%。珊瑚礁鱼类共14科28属36种, 其中, 雀鲷科的种类数最多, 为11种。鹿回头4 m断面珊瑚礁鱼类密度最大, 为154尾/300 m²。砗磲和龙虾极少发现, 珊瑚天敌核果螺多见。东岛、鹿回头、大东海珊瑚礁生态系统健康状况均处于“一般”。本文所采用的方法是结合常规珊瑚礁监测可获得的指标进行评价, 简便易操作, 通过在三亚珊瑚礁保护区的实践, 能够很好地反映珊瑚礁生态系统现状及其健康状况, 科研和业务化监测部门均可应用。     

Historical and contemporary determinants of global phylogenetic structure in tropical reef fish faunas

Identifying the main determinants of tropical marine biodiversity is essential for devising appropriate conservation measures mitigating the ongoing degradation of coral reef habitats. Based on a gridded distribution database and phylogenetic information, we compared the phylogenetic structure of assemblages for three tropical reef fish families (Labridae: wrasses, Pomacentridae: damselfishes and Chaetodontidae: butterflyfishes) using the net relatedness (NRI) and nearest taxon (NTI) indices. We then related these indices to contemporary and historical environmental conditions of coral reefs using spatial regression analyses. Higher levels of phylogenetic clustering were found for fish assemblages in the Indo‐Australian Archipelago (IAA), and more particularly when considering the NTI index. The phylogenetic structure of the Pomacentridae, and to a lower extent of the Chaeotodontidae and Labridae, was primarily associated with the location of refugia during the Quaternary period. Phylogenetic clustering in the IAA may partly result from vicariance events associated with coral reef fragmentation during the glacial periods of the Quaternary. Variation in the patterns among fish families further suggest that dispersal abilities may have interacted with past habitat availability in shaping the phylogenetic structure of tropical reef fish assemblages.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Growth of reef fishes in response to live coral cover

Although the global decline in coral reef health is likely to have profound effects on reef associated fishes, these effects are poorly understood. While declining coral cover can reduce the abundance of reef fishes through direct effects on recruitment and/or mortality, recent evidence suggests that individuals may survive in disturbed habitats, but may experience sublethal reductions in their condition. This study examined the response of 2 coral associated damselfishes (Pomacentridae), Chrysiptera parasema and Dascyllus melanurus, to varying levels of live coral cover. Growth, persistence, and the condition of individuals were quantified on replicate coral colonies in 3 coral treatments: 100% live coral (control), 50% live coral (partial) and 0% live coral (dead). The growth rates of both species were directly related to the percentage live coral cover, with individuals associated with dead corals exhibiting the slowest growth, and highest growth on control corals. Such differences in individual growth between treatments were apparent after 29 d. There was no significant difference in the numbers of fishes persisting or the physiological condition of individuals between different treatments on this time-scale. Slower growth in disturbed habitats will delay the onset of maturity, reduce lifetime fecundity and increase individual's vulnerability to gape-limited predation. Hence, immediate effects on recruitment and survival may underestimate the longer-term impacts of declining coral on the structure and diversity of coral-associated reef fish communities.     

Patterns of morphological evolution of the cephalic region in damselfishes (Perciformes: Pomacentridae) of the Eastern Pacific

Pomacentridae are one of the most abundant fish families inhabiting reefs of tropical and temperate regions. This family, comprising 29 genera, shows a remarkable diversity of habitat preferences, feeding, and behaviours. Twenty‐four species belonging to seven genera have been reported in the Eastern Pacific region. The present study focuses on the relationship between the diet and the cephalic profile in the 24 endemic damselfishes of this region. Feeding habits were determined by means of underwater observations and the gathering of bibliographic data. Variations in cephalic profile were analyzed by means of geometric morphometrics and phylogenetic methods. The present study shows that the 24 species can be grouped into three main trophic guilds: zooplanktivores, algivores, and an intermediate group feeding on small pelagic and benthic preys. Shape variations were low within each genus except for Abudefduf. Phylogenetically adjusted regression reveals that head shape can be explained by differences in feeding habits. The morphometric phylogeny recovered the subfamily Stegastinae and the relationship between Abudefduf troschelii and Chromis species. The cephalic profile of damselfishes contains a clear and strong phylogenetic signal. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 593–613.     

Molecular phylogenetic evidence for the evolution of specialization in anemonefishes.

Anemonefishes (genera: Amphiprion and Premnas; family Pomacentridae) are a group of 28 species of coral reef fishes that are found in obligate symbiosis with large tropical sea anemones. A phylogenetic hypothesis based on morphological analyses of this group suggests that the ancestral anemonefish was a generalist with similar morphology to other pomacentrids, and that it gave rise to other anemonefish species that were more specialized for living with particular species of host anemones. To test this hypothesis we constructed a molecular phylogeny for the anemonefishes by sequencing 1140 base pairs of the cytochrome b gene and 522 base pairs of the 16S rRNA gene for six species of anemonefishes (representatives of all subgenera and species complexes) and two other pomacentrid species. Three methods of phylogenetic analysis all strongly supported the conclusion that anemonefishes are a monophyletic group. The molecular phylogeny differs from the tree based on morphological data in that the two species of specialized anemonefishes (Premnas biaculeatus and Amphiprion ocellaris) were assigned to a basal position within the clade, and the extreme host generalist (Amphiprion clarkii) to a more derived position. Thus, the initial anemonefish ancestors were probably host specialists and subsequent speciation events led to a combination of generalist and specialist groups. Further phylogenetic studies of additional anemonefish species are required to substantiate this hypothesis.     

Consumption of coral propagules represents a significant trophic link between corals and reef fish

Mass spawning of corals provides a large seasonal pulse of high-energy prey that potentially benefits reef fish that are capable of capturing and digesting coral propagules. This study examines the range of fish species that consume coral propagules and also tests whether reef fish experience a significant increase in physiological condition when feeding on coral propagules. Thirty-six species of diurnal reef fish were seen to consume coral propagules released during mass coral spawning. Stomach content analyses of three reef fish species (Pomacentrus moluccensis, Abudefduf whitleyi, and Caesio cunning) revealed that both P. moluccensis and A. whitleyi feed almost exclusively on coral propagules during mass coral spawning. Fish feeding extensively on coral propagules also amassed considerable lipid stores, which could greatly improve the quality and survivorship of their progeny. In contrast, C. cunning consumed only very small quantities of coral propagules, and showed no detectable change in lipid stores during the course of the study. This study provides the first direct evidence that reef fish benefit from mass coral spawning, and reveals a potentially significant trophic link between scleractinian corals and reef fish. Accepted: 9 June 2000     

Increasing ocean temperature reduces the metabolic performance and swimming ability of coral reef damselfishes

Tropical coral reef teleosts are exclusively ectotherms and their capacity for physical and physiological performance is therefore directly influenced by ambient temperature. This study examined the effect of increased water temperature to 3 °C above ambient on the swimming and metabolic performance of 10 species of damselfishes (Pomacentridae) representing evolutionary lineages from two subfamilies and four genera. Five distinct performance measures were tested: (a) maximum swimming speed (U_crit), (b) gait‐transition speed (the speed at which they change from strictly pectoral to pectoral‐and‐caudal swimming, U_p?c), (c) maximum aerobic metabolic rate (MO_2?MAX), (d) resting metabolic rate (MO_2?REST), and (e) aerobic scope (ratio of MO_2?MAX to MO_2?REST, A_SC). Relative to the control (29 °C), increased temperature (32 °C) had a significant negative effect across all performance measures examined, with the magnitude of the effect varying greatly among closely related species and genera. Specifically, five species spanning three genera (Dascyllus, Neopomacentrus and Pomacentrus) showed severe reductions in swimming performance with U_crit reduced in these species by 21.3–27.9% and U_p?c by 32.6–51.3%. Furthermore, five species spanning all four genera showed significant reductions in metabolic performance with aerobic scope reduced by 24.3–64.9%. Comparisons of remaining performance capacities with field conditions indicate that 32 °C water temperatures will leave multiple species with less swimming capacity than required to overcome the water flows commonly found in their respective coral reef habitats. Consequently, unless adaptation is possible, significant loss of species may occur if ocean warming of ≥3 °C arises.     

Temperature influences habitat preference of coral reef fishes: Will generalists become more specialised in a warming ocean?

Climate change is expected to pose a significant risk to species that exhibit strong behavioural preferences for specific habitat types, with generalist species assumed to be less vulnerable. In this study, we conducted habitat choice experiments to determine how water temperature influences habitat preference for three common species of coral reef damselfish (Pomacentridae) that differ in their levels of habitat specialisation. The lemon damselfish Pomacentrus moluccensis, a habitat specialist, consistently selected complex coral habitat across all temperature treatments (selected based on local average seasonal temperatures naturally experienced in situ: ambient winter 22°C; ambient summer 28°C; and elevated 31°C). Unexpectedly, the neon damselfish Pomacentrus coelestis and scissortail sergeant Abudefduf sexfasciatus, both of which have more generalist habitat associations, developed strong habitat preferences (for complex coral and boulder habitat, respectively) at the elevated temperature treatment (31°C) compared to no single preferred habitat at 22°C or 28°C. The observed shifts in habitat preference with temperature suggest that we may be currently underestimating the vulnerability of some habitat generalists to climate change and highlight that the ongoing loss of complex live coral through coral bleaching could further exacerbate resource overlap and species competition in ways not currently considered in climate change models.     

Evolution of a Neotropical marine fish lineage (Subfamily Chaenopsinae, Suborder Blennioidei) based on phylogenetic analysis of combined molecular and morphological data

Phylogenetic relationships within tube blennies (Chaenopsinae) were reconstructed using Bayesian, maximum parsimony and likelihood analyses of multiple molecular markers (mitochondrial DNA: COI; nuclear DNA: TMO-4C4, RAG1, Rhodopsin, and Histone H3) and 148 morphological characters. This total-evidence based topology is well-resolved and congruent across analytical methods with strong support for the monophyly of the Chaenopsinae, all included genera and several internal nodes. A rapid radiation in the early evolution of chaenopsins is inferred from the relatively poor support values for relationships among basal lineages and their divergence into different habitats (rocky reefs, coral reefs and the reef/sand interface). Rates of molecular evolution in chaenopsins, as inferred by divergence among four putative transisthmian geminate species pairs, are rapid compared to other fishes. Conflicts among genetic markers and morphology are especially evident within the genus Coralliozetus, with different species relationships supported by morphology, TMO-4C4, and RAG1 plus Rhodopsin. This study hypothesizes a novel sistergroup relationship between Ekemblemaria and Hemiemblemaria, consistent with morphological, molecular and habitat use data. Our total evidence phylogenetic hypothesis indicates that previously hypothesized morphological characters supporting a close relationship between Hemiemblemaria and Chaenopsis plus Lucayablennius resulted from convergent evolution in these relatively free-swimming blennies.     

Experimental artificial cultivation of corals from colony fragments

In 2010–2011, experimental commercial cultivation of 14 species of hermatypic corals was carried out using the method of donor colony fragmentation. The transplants successfully survived on experimental frame installations. The coral colonies that were recovered from the fragments became attached to the frame installations in a similar way to their attachment on natural substrata. After 1 year, the size of the trans-plants was found to have increased by 275%. The newly formed artificial coral community was colonized by the damselfish Dascyllus reticulates (Pomacentridae); the species is a common coral fish species that lives on natural reefs.     

Species recognition in the blackbordered damselfish Dascyllus marginatus (Rüppell): An evaluation of computer-animated playback techniques

Blackbordered damselfish, Dascyllus marginatus, were presented with computer animations of varying backgrounds, with and without moving stimuli of conspecifics and sympatric heterospecifics. When subjects were released into the test tank their initial escape response was nearly always towards a fish stimulus, independent of species identity, rather than towards a background stimulus, even when the former where presented in the open and the latter displayed a coral shelter. Subjects did, however, show distinct preferences according to species with regards to the total time spent with each animation. D. marginatus significantly preferred conspecifics over Dascyllus trimaculatus, but did not prefer conspecifics over Dascyllus aruanus, a species with which they frequently shoal in nature. These results demonstrate that computer animations can be used in ethological studies of coral reef fishes and to isolate critical visual signals. Results also show that reef fishes can use visual signals to make sophisticated discriminations among species that can be modulated according to the task at hand.     

Planktonic and symbiotic organisms in nutrition of coralobiont fish

Nutrition of coral reef fish closely associated with the colonies of scleractinian corals Acropora spp. (South China Sea, Gulf of Nha Trang) was investigated. Parabionts Dascyllus reticulatus and Chromis caeruleus (Pomacentridae) feed mainly on zooplankton in the water column above the hosts’ colony, and inbionts Gobiodon quinquestrgatus (Gobiidae) rarely leave a host colony and feed generally on coral reef invertebrates. A high part of secondary nutrition objects (on average 19–29%) that indicates deficiency in main food were noted in both categories of fish. Food of all three fish species contains parasites (from 19% in C. caeruleus and up to 25% in G. quinquestrigatus) whose significant part is represented by parasites of coral (approximately 7% in D. reticulatus and C. caeruleus and 15% in G. quinquestrigatus). In addition, larvae of parasites of Isopoda and Facetotecta that are fed by fish in the water column were found. Mucus from the surface of host coral which was recognized on the basis of a high number of nematocysts in fish stomachs was found in food of G. quinquestrigatus for the first time. Positive and negative effects of symbiosis of fish and host-coral were discussed. A commensal character of symbiosis was supposed.     

Auditory sensitivity in settlement-stage larvae of coral reef fishes

The larval phase of most species of coral reef fishes is spent away from the reef in the pelagic environment. At the time of settlement, these larvae need to locate a reef, and recent research indicates that sound emanating from reefs may act as a cue to guide them. Here, the auditory abilities of settlement-stage larvae of four species of coral reef fishes (families Pomacentridae, Lutjanidae and Serranidae) and similar-sized individuals of two pelagic species (Carangidae) were tested using an electrophysiological technique, auditory brainstem response (ABR). Five of the six species heard frequencies in the 100–2,000 Hz range, whilst one carangid species did not detect frequencies higher than 800 Hz. The audiograms of the six species were of similar shape, with best hearing at lower frequencies between 100 and 300 Hz. Strong within-species differences were found in hearing sensitivity both among the coral reef species and among the pelagic species. Larvae of the coral reef species had significantly more sensitive hearing than the larvae of the pelagic species. The results suggest that settlement-stage larval reef fishes may be able to detect reef sounds at distances of a few 100 m. If true hearing thresholds are lower than ABR estimates, as indicated in some comparisons of ABR and behavioural methods, the detection distances would be much larger.     

Molecular systematics of flyingfishes (Teleostei: Exocoetidae): evolution in the epipelagic zone

The flyingfish family Exocoetidae is a diverse group of marine fishes that are widespread and abundant in tropical and subtropical seas. Flyingfishes are epipelagic specialists that are easily distinguished by their enlarged fins, which are used for gliding leaps over the surface of the water. Although phylogenetic hypotheses have been proposed for flyingfish genera based on morphology, no comprehensive molecular studies have been performed. In the present study, we describe a species‐level molecular phylogeny for the family Exocoetidae, based on data from the mitochondrial cytochrome b gene (1137 bp) and the nuclear RAG2 gene (882 bp). We find strong support for previous morphology‐based phylogenetic hypotheses, as well as the monophyly of most currently accepted flyingfish genera. However, the most diverse genus Cheilopogon is not monophyletic. Using our novel flyingfish topology, we examine previously proposed hypotheses for the origin and evolution of gliding. The results support the progressive transition from two‐wing to four‐wing gliding. We also use phylogenetic approaches to test the macroecological effects of two life history characters (e.g. egg buoyancy and habitat) on species range size in flyingfishes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 161–174.     

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.