Life history of the symbiotically luminous cardinalfish Siphamia tubifer (Perciformes: Apogonidae)

Characteristics of the life history of the coral reef‐dwelling cardinalfish Siphamia tubifer, from Okinawa, Japan, were defined. A paternal mouthbrooder, S. tubifer, is unusual in forming a bioluminescent symbiosis with Photobacterium mandapamensis. The examined S. tubifer (n = 1273) ranged in size from 9·5 to 43·5 mm standard length (L_S), and the minimum size at sexual maturity was 22 mm L_S. The number of S. tubifer associated during the day among the spines of host urchins was 22·9 ± 16·1 (mean ± s.d .; Diadema setosum) and 3·6 ± 3·2 (Echinothrix calamaris). Diet consisted primarily of crustacean zooplankton. Batch fecundity (number of eggs; F_B) was related to L_S by the equations: males (fertilized eggs) F_B = 27·5L_S ? 189·46; females (eggs) F_B = 31·3L_S ? 392·63. Individual mass (M; g) as a function of L_S was described by the equation: . Growth, determined from otolith microstructure analysis, was described with the von Bertalanffy growth function with the following coefficients: L_∞ = 40·8 mm L_S, K = 0·026 day^?1 and t₀ = 23·25 days. Planktonic larval duration was estimated to be 30 days. The age of the oldest examined individual was 240 days. The light organ of S. tubifer, which harbours the symbiotic population of P. mandapamensis, increased linearly in diameter as S. tubifer L_S increased, and the bacterial population increased logarithmically with S. tubifer L_S. These characteristics indicate that once settled, S. tubifer grows quickly, reproduces early and typically survives much less than 1 year in Okinawa. These characteristics are generally similar to other small reef fishes but they indicate that S. tubifer experiences higher mortality.     

Symbiosis initiation in the bacterially luminous sea urchin cardinalfish Siphamia versicolor

To determine how each new generation of the sea urchin cardinalfish Siphamia versicolor acquires the symbiotic luminous bacterium Photobacterium mandapamensis, and when in its development the S. versicolor initiates the symbiosis, procedures were established for rearing S. versicolor larvae in an aposymbiotic state. Under the conditions provided, larvae survived and developed for 28 days after their release from the mouths of males. Notochord flexion began at 8 days post release (dpr). By 28 dpr, squamation was evident and the caudal complex was complete. The light organ remained free of bacteria but increased in size and complexity during development of the larvae. Thus, aposymbiotic larvae of the fish can survive and develop for extended periods, major components of the luminescence system develop in the absence of the bacteria and the bacteria are not acquired directly from a parent, via the egg or during mouth brooding. Presentation of the symbiotic bacteria to aposymbiotic larvae at 8-10 dpr, but not earlier, led to initiation of the symbiosis. Upon colonization of the light organ, the bacterial population increased rapidly and cells forming the light-organ chambers exhibited a differentiated appearance. Therefore, the light organ apparently first becomes receptive to colonization after 1 week post-release development, the symbiosis is initiated by bacteria acquired from the environment and bacterial colonization induces morphological changes in the nascent light organ. The abilities to culture larvae of S. versicolor for extended periods and to initiate the symbiosis in aposymbiotic larvae are key steps in establishing the experimental tractability of this highly specific vertebrate and microbe mutualism.     

LuxA gene of light organ symbionts of the bioluminescent fish Acropoma japonicum (Acropomatidae) and Siphamia versicolor (Apogonidae) forms a lineage closely related to that of Photobacterium leiognathi ssp. mandapamensis

A molecular phylogenetic analysis of luxA gene sequences of light organ symbionts of the fish Acropoma japonicum (Acropomatidae) and Siphamia versicolor (Apogonidae) revealed that the sequences were related to those of Photobacterium leiognathi ssp. mandapamensis, which is not known to occur as a light organ symbiont among bioluminescent P. leiognathi clades. The presence of another lux gene element, luxF, coding for nonfluorescent protein, provided additional support for the identity of the light organ symbionts of the fish. Cladogenesis of the light organ symbiont P. leiognathi may be influenced by the radiation of host fishes.     

Comparative oogenesis in cardinal fishes (Apogonidae,Perciformes), with special focus on the adaptive structures of the egg envelopes

The present study compares gonad structure and oogenesis in 30 species of cardinal fish (Apogonidae) from the Indo-Pacific region. In all of the species studied the ovaries are bilobed, with each lobe possessing ovigerous lamellae whose numbers and dimensions increase concomitantly with growth of the fish and reproductive stage, and correlate with the total dimensions of the particular species. In species of the genus Siphamia, the ovarian lobes are associated with a fat body that is not found in species of the other genera studied. Some of the species studied (Siphamia spp.; Archamia spp.) demonstrate group-synchronized maturation of eggs, possibly with only one spawn per reproduction season. However, most of the Apogon spp., and other genera, have unsynchronized cycles, a characteristic of multiple spawners. The various stages of previtellogenic and vitellogenic development are similar in the different species, but differ in timing and dimensions of the oocytes around which the morphogenesis of the egg envelope (chorion) and follicle begins. Differentiation of the egg envelope and follicle cells parallels the vitellogenic stages of egg development. The number of eggs was found to correlate positively with the length of the body and ovary of the fish, whereas egg diameter was found to correlate negatively with body length. The surface of the egg envelope bears ridges that form patterns of various types, whose structures and dimensions seem to be species-specific, and which in most species converge upon the single micropyle on the animal pole. The exposed ridge mazes include a special fibrous web that anchors in the micropyle. The production and importance of these ridges, as well as the correlations between egg numbers per spawn, egg dimensions, and the oral dimensions of the paternal male, are discussed.     

Long-term cleaner fish presence affects growth of a coral reef fish

Cleaning behaviour is considered to be a classical example of mutualism. However, no studies, to our knowledge, have measured the benefits to clients in terms of growth. In the longest experimental study of its kind, over an 8 year period, cleaner fish Labroides dimidiatus were consistently removed from seven patch reefs (61-285 m(2)) and left undisturbed on nine control reefs, and the growth and parasite load of the damselfish Pomacentrus moluccensis determined. After 8 years, growth was reduced and parasitic copepod abundance was higher on fish from removal reefs compared with controls, but only in larger individuals. Behavioural observations revealed that P. moluccensis cleaned by L. dimidiatus were 27 per cent larger than nearby conspecifics. The selective cleaning by L. dimidiatus probably explains why only larger P. moluccensis individuals benefited from cleaning. This is the first demonstration, to our knowledge, that cleaners affect the growth rate of client individuals; a greater size for a given age should result in increased fecundity at a given time. The effect of the removal of so few small fish on the size of another fish species is unprecedented on coral reefs.     

Apogon smithi (Kotthaus, 1970) (Teleostei: Apogonidae), a Red Sea cardinalfish colonizing the Mediterranean Sea

Smith's cardinalfish Apogon smithi is recorded for the first time in the Mediterranean Sea. Six specimens were caught by commercial trawler north of Jaffa, Israel, at depths of 40 m and the relatively high number of specimens suggests that A. smithi has established a sustainable population. This is the third Indo-Pacific apogonid species documented in the Mediterranean Sea, and is assumed to be a migrant originating from the Red Sea.     

A new species of the cardinalfish genus Gymnapogon (Perciformes, Apogonidae) from the Red Sea

 A new cardinalfish species, Gymnapogon melanogaster, is described from two specimens collected at night in the Gulf of Aqaba, Eilat, Israel. This species is characterized by having 9 dorsal and 8 anal fin soft rays; 14–15 pectoral fin rays; 2 + 11 gill rakers; a flat, bifurcated preopercular spine; a naked body without a papillae network; black pelvic fins; and a black stomach. It is similar to Gymnapogon vanderbilti (Fowler, 1938) that is known only from the Line Islands of the Central Pacific Ocean. Received: December 26, 2001 / Revised: June 10, 2002 / Accepted: June 24, 2002 Acknowledgments We thank D. Didier and M. Sabaj of the Academy of Natural Sciences, Philadelphia, for loans of and for taking data from type specimens; T.H. Fraser of the Mote Marine Laboratory, Sarasota, kindly provided data on type specimens. We are grateful to E. Heemstra of the South African Institute for Aquatic Biodiversity, Grahamstown, South Africa, for the artwork presented in this article and to A. Lerner of the Hebrew University, Jerusalem, for his assistance in collecting the specimens. Correspondence to:Ofer Gon     

Functional significance of genetically different symbiotic algae Symbiodinium in a coral reef symbiosis

The giant sea anemone Condylactis gigantea associates with members of two clades of the dinoflagellate alga Symbiodinium, either singly or in mixed infection, as revealed by clade-specific quantitative polymerase chain reaction of large subunit ribosomal DNA. To explore the functional significance of this molecular variation, the fate of photosynthetically fixed carbon was investigated by (14)C radiotracer experiments. Symbioses with algae of clades A and B released ca. 30-40% of fixed carbon to the animal tissues. Incorporation into the lipid fraction and the low molecular weight fraction dominated by amino acids was significantly higher in symbioses with algae of clade A than of clade B, suggesting that the genetically different algae in C. gigantea are not functionally equivalent. Symbioses with mixed infections yielded intermediate values, such that this functional trait of the symbiosis can be predicted from the traits of the contributing algae. Coral and sea anemone symbioses with Symbiodinium break down at elevated temperature, a process known as 'coral bleaching'. The functional response of the C. gigantea symbiosis to heat stress varied between the algae of clades A and B, with particularly depressed incorporation of photosynthetic carbon into lipid of the clade B algae, which are more susceptible to high temperature than the algae of clade A. This study provides a first exploration of how the core symbiotic function of photosynthate transfer to the host varies with the genotype of Symbiodinium, an algal symbiont which underpins corals and, hence, coral reef ecosystems.     

First record of Apogon queketti Gilchrist (Osteichthyes: Apogonidae) in the Mediterranean Sea

The spotfin cardinalfish Apogon queketti is recorded for the first time in the north-eastern Mediterranean Sea (Iskenderun Bay, Turkey). Two specimens of this species, 111 and 102 mm total length were caught by a commercial trawler at depths of 55 and 60 m on 14 October 2004 and 23 April 2005, respectively. The occurrence of this species in the Mediterranean Sea is the result of migration from the Red Sea via the Suez Canal.     

Spatial structure of coral reef fish communities at Kudaka Island (Ryukyu Archipelago), Japan

The present study aimed to investigate the spatial structure of fish communities at juvenile and adult stages on coral reefs at Kudaka Island (Ryukyu Archipelago, Japan) and to relate spatial patterns in the structure of the fish communities to gradients in environmental variables. Diurnal visual censuses allowed us to record 2,602 juveniles belonging to 60 species and 1,543 adults belonging to 53 species from October to December 2005. The distribution of species highlighted that the juvenile community was organised into three distinct assemblages, rather than exhibiting gradual change in community structure along the cross-reef gradient. Correlations between spatial patterns of juvenile community and environmental variables revealed that the most significant factors explaining variation in community structure were coral rubble and coral slab. In contrast, the adult community was organised into one assemblage, and the most significant variation factors in community structure were depth, live coral in massive form, live coral in branched form, dead coral and sand. Overall, the present study showed that most juvenile and adult coral reef fish at Kudaka Island exhibited striking patterns in their distribution and depth and some biological factors (e.g., abundance of live coral, dead coral and coral rubble) might exert considerable influence on the distribution of fishes.     

Traits,landmarks and outlines: Three congruent sides of a tale on coral reef fish morphology

Quantifying the morphology of organisms remains fundamental in ecology given the form‐function relationship. Morphology is quantifiable in traits, landmarks, and outlines, and the choice of approach may influence ecological conclusions to an unknown extent. Here, we apply these three approaches to 111 individual coral reef fish of 40 species common in Micronesia. We investigate the major dimensions of morphological variability among individuals, families, and predefined feeding functional groups. We find that although the approaches are complementary, they coincide in capturing elongation as the main dimension of variability. Furthermore, the choice of approach led to different interpretations regarding the degree of morphological differentiation among taxonomic and feeding functional groups. We also use each morphology dataset to compute community‐scale morphological diversity on Palauan reefs and investigate how the choice of dataset affects the detection of differences among sites and wave exposure levels. The exact ranking of sites from highest to lowest morphological diversity was sensitive to the approach used, but not the broad spatial pattern of morphological diversity. Conclusions regarding the effect of wave exposure on morphological diversity were robust to the approach used. Biodiversity hotspots (e.g., areas of exceptionally high diversity and/or endemism) are considered important conservation targets but their location may depend on the biodiversity metric used. In the same vein, our results caution against labelling particular sites as morphological diversity hotspots when metrics consider only a single aspect of morphology.     

Exploring the nature of ecological specialization in a coral reef fish community: morphology,diet and foraging microhabitat use

Patterns of ecological specialization offer invaluable information about ecosystems. Yet, specialization is rarely quantified across several ecological niche axes and variables beyond the link between morphological and dietary specialization have received little attention. Here, we provide a quantitative evaluation of ecological specialization in a coral reef fish assemblage (f. Acanthuridae) along one fundamental and two realized niche axes. Specifically, we examined ecological specialization in 10 surgeonfish species with regards to morphology and two realized niche axes associated with diet and foraging microhabitat utilization using a recently developed multidimensional framework. We then investigated the potential relationships between morphological and behavioural specialization. These relationships differed markedly from the traditional ecomorphological paradigm. While morphological specialization showed no relationship with dietary specialization, it exhibited a strong relationship with foraging microhabitat specialization. However, this relationship was inverted: species with specialized morphologies were microhabitat generalists, whereas generalized morphotypes were microhabitat specialists. Interestingly, this mirrors relationships found in plant–pollinator communities and may also be applicable to other ecosystems, highlighting the potential importance of including niche axes beyond dietary specialization into ecomorphological frameworks. On coral reefs, it appears that morphotypes commonly perceived as most generalized may, in fact, be specialized in exploiting flat and easily accessible microhabitats.     

Paranthias furcifer (Perciformes: Serranidae), a new alien fish in the Mediterranean Sea

A specimen of a creole fish Paranthias furcifer (285 mm total length) was captured in Marina Bay (Croatian coast) in the eastern Adriatic Sea. This is the first Mediterranean record of this species. The possible modes of introduction of species are discussed.     

Life-history traits of a widely distributed coral reef fish,Mulloidichthys flavolineatus (Mullidae), Okinawa Island,southern Japan

Environmental Biology of Fishes - Yellowstripe goatfish, Mulloidichthys flavolineatus, is a widely distributed commercial fish in the Indo-Pacific. However, there have been few comprehensive...     

Functional role of the fractal morphology of corals: A full model of the nutrient turbulent diffusion fluxes to a coral reef

We present for the first time a dynamic model relating the functional role played by a fractal morphology: the effect of the fractal texture of adsorbents on the turbulent diffusion flux enhancement is quantified. The resulting law is extrapolated to a set of fractal dimensions of coral colonies. Thus, we reach the conclusion that previous modelisation of a reef nutrient uptake did not match with field values, because it only took the colony scale roughness into account, whereas our multiscale fractal approach matches real data very well.     

Evolutionary history of the reef fish Anisotremus interruptus (Perciformes: Haemulidae) throughout the Tropical Eastern Pacific

The Tropical Eastern Pacific (TEP) is a dynamic coastal environment characterized by a complex system of oceanic processes and discontinuous rocky habitats. These features, in conjunction with the ecological and physiological characteristics of Anisotremus interruptus, might limit gene flow and shape the evolutionary history of the species. In this study, we investigate the evolutionary history of the reef fish A. interruptus (and its Atlantic sister species A. surinamensis) throughout its range in the TEP, using two mitochondrial (cox1 and cytb) and two nuclear markers (S7 and RAG1). We found three genetic groups of A. interruptus with recent divergence times from the Galapagos Archipelago, Revillagigedo Archipelago, the continental TEP, and A. surinamensis the sister specie from the Atlantic. The haplotype mtDNA networks show A. surinamensis in a central position with respect to Pacific genetic haplogroups, whereas nDNA networks show mixed haplotypes between the four genetic groups. In the species tree, A. surinamensis appears as the sister species of all the Pacific samples and the Galapagos Archipelago population emerges as a genetically distinctive group. The samples from the Revillagigedo Archipelago also constitute a genetic distinctive group, closely related to the continental samples. Continental individuals do not show significant genetic structure and exhibit a population expansion during the Pleistocene. The sandy gaps of the TEP not appear to act as barriers isolating populations of A. interruptus, whereas the open sea gap between the oceanic islands and the continental coast do.     

Habitat complexity modifies the impact of piscivores on a coral reef fish population

Patterns in juvenile mortality rates can have a profound affect on the distribution and abundance of adult individuals, and may be the result of a number of interacting factors. Field observations at Lizard Island (Great Barrier Reef, Australia) showed that for a coral reef damselfish, Pomacentrus moluccensis, juvenile mortality (over 1 year) varied between 20 and almost 100% among sites. Correlative data showed that juvenile mortality increased as a function of initial densities (recruitment), predator densities and the availability of preferred coral substrata. A multiple regression showed that these three variables together did not explain significantly more variation in mortality than the single factor showing the strongest relationship. This appeared to be because recruitment, predator densities and preferred coral substrata were all highly correlated, suggesting that one, two or all of these factors may be influencing juvenile mortality rates. One hypothesis was that density-dependent mortality in juveniles was the result of an interaction between predators (which appear to aggregate at high-recruitment sites) and the availability of preferred substrata (predator refuges). We tested this hypothesis by using both laboratory and field experiments to see whether fish predation could significantly alter survivorship of this damselfish, and whether this impact was dependent upon the coral substratum. The laboratory experiment was designed to test the effects of three common predators (Pseudochromis fuscus, Cephalopholis boenak and Thalassoma lunare) and three different coral substrata that varied in their complexity (Pocillopora damicornis, Acropora nasuta and A. nobilis) on the survival of juvenile Pomacentrus moluccensis. There was a significant interaction between predator species and microhabitat in determining survival. Pseudochromis fuscus and C. boenak were both significantly better at capturing juvenile damselfish than T. lunare. Juvenile survivorship was significantly better when they were given the more complex corals, Pocillopora damicornis and A. nasuta, compared with those given the open-structured species A. nobilis. This pattern reflects habitat selection in the field. Predators differed in their strike rates and the proportion of strikes that were successful, but all exhibited greater success at prey capture where A. nobilis was provided as shelter. The interaction between the effect of predator species and microhabitat structure on damselfish survival was tested in the field for a cohort of juvenile Pomacentrus moluccensis. We examined juvenile survival in the presence and absence of two predators that co-occur on natural patch reefs (C. boenak and Pseudochromis fuscus). The experimental patch reefs we used for this purpose were constructed from both high complexity (Pocillopora damicornis) and low complexity (A. nobilis) coral substrata. Both juveniles and predators were translocated to reefs at natural densities. The effects of predation were clearly dependent upon the microhabitat. Reefs of the high-complexity coral with predators supported the same high numbers of Pomacentrus moluccensis as the reefs with no resident predators. However, damselfish abundance was significantly lower on low-complexity reefs with resident predators, relative to the other treatments. Background rates of loss were high, even on preferred coral in the absence of the manipulated predator, suggesting that transient predators may be even more important than the residents. We suggest that adult abundances in this species were strongly influenced by the densities of different predators and the availability of preferred refuges. Received: 3 April 1997 / Accepted: 26 August 1997