Nocturnal substrate association of four coral reef fish groups (parrotfishes, surgeonfishes, groupers and butterflyfishes) in relation to substrate architectural characteristics.

Although numerous coral reef fish species utilize substrates with high structural complexities as habitats and refuge spaces, quantitative analysis of nocturnal fish substrate associations has not been sufficiently examined yet. The aims of the present study were to clarify the nocturnal substrate associations of 17 coral reef fish species (nine parrotfish, two surgeonfish, two grouper and four butterflyfish) in relation to substrate architectural characteristics. Substrate architectural characteristics were categorized into seven types: (1) eave-like space, (2) large inter-branch space, (3) overhang by protrusion of fine branching structure, (4) overhang by coarse structure, (5) uneven structure without large space or overhang, (6) flat and (7) macroalgae. Overall, fishes were primarily associated with three architectural characteristics (eave-like space, large inter-branch space and overhang by coarse structure). The main providers of these three architectural characteristics were tabular and corymbose Acropora, staghorn Acropora, and rock. Species-specific significant positive associations with particular architectural characteristics were found as follows. For the nine parrotfish species, Chlorurus microrhinos with large inter-branch space and overhang by coarse structure; Ch. spilurus with eave-like space and large inter-branch space; Hipposcarus longiceps with large inter-branch space; Scarus ghobban with overhang by coarse structure; five species (Scarus forsteni, S. niger, S. oviceps, S. rivulatus and S. schlegeli) with eave-like space. For the two surgeonfish species, Naso unicornis with overhang by coarse structure; N. lituratus with eave-like space. For the two grouper species, Plectropomus leopardus with eave-like space; Epinephelus ongus with overhang by coarse structure. For the four butterflyfish species, Chaetodon trifascialis with eave-like space and large inter-branch space; C. lunulatus and C. ephippium with large inter-branch space; C. auriga showed no significant associations with any architectural characteristics. Four species (Ch. microrhinos, H. longiceps, S. niger and N. unicornis) also showed clear variations in substrate associations among the different fish size classes. Since parrotfishes, surgeonfishes and groupers are main fisheries targets in coral reefs, conservation and restoration of coral species that provide eave-like space (tabular and corymbose Acropora) and large inter-branch space (staghorn Acropora) as well as hard substrates with coarse structure that provide overhang (rock) should be considered for effective fisheries management in coral reefs. For butterflyfishes, coral species that provide eave-like space (tabular Acropora) and large inter-branch space (staghorn Acropora) should also be conserved and restored for provision of sleeping sites.

Spawning aggregations of checkered snapper (Lutjanus decussatus) and blackspot snapper (L. fulviflamma): seasonality, lunar-phase periodicity and spatial distribution within spawning ground.

Snappers (family Lutjanidae) are important fisheries target species and some species are known to form spawning aggregations at particular spawning grounds. The present study investigated the ecological characteristics of fish aggregations of two snapper species (checkered snapper Lutjanus decussatus and blackspot snapper L. fulviflamma) that form at a particular site. Specifically, the aims were to clarify (1) seasonality and lunar-phase periodicity of fish aggregation formation, (2) fine-scale spatial distribution of fish density (spatial variations of fish density at intervals of several-tens meters) within the aggregation site, (3) size and age frequency distributions of fishes in the aggregation site, (4) gonad development, (5) to compare fish abundance between inside and outside the aggregation site, and (6) to verify that fish aggregations of the two snapper species were spawning aggregation. Underwater observations using a 600 m × 5 m transect revealed that greater fish abundance of Lutjanus decussatus was found monthly between May and October, and clear positive peaks in the fish abundance were found only around the last-quarter moon. This lunar-related periodicity in the increase of fish abundance was confirmed by a time-series analysis (correlogram). Within the aggregation site, L. decussatus showed a relatively uniform distribution. In contrast, greater fish abundance of L. fulviflamma was found monthly between April and October, and clear positive peaks in the fish abundance were found around the last-quarter moon (April, May, June and October) or new moon (July, August and September). This lunar-related periodicity was also confirmed by correlogram. Lutjanus fulviflamma showed a relatively clumped distribution within the aggregation site. Most females of the two species in the aggregation site had hydrated eggs, indicating that the two species form aggregations for reproduction. The two species, although occurring simultaneously, are considered to form aggregations of conspecifics only. For L. decussatus, average fork length and age of males and females were 229.2 mm and 243.9 mm and 9.4 years and 8.1 years, respectively. For L. fulviflamma, average fork length and age of males and females were 233.9 mm and 246.9 mm and 6.8 years and 8.1 years, respectively. Fish abundance inside the aggregation site was 266.8-fold and 141557.1-fold greater than those outside the aggregation site for L. decussatus and L. fulviflamma, respectively. These results showed that (1) fish aggregation formation of the two snapper species was predictably repeated in particular months and lunar-phase, (2) it was predictably found at the particular site, (3) the fish abundance in the aggregation site markedly exceeded the fish abundance outside the aggregation site, and (4) the two species form aggregations for reproduction. Therefore, it is suggested that the fish aggregations for the two species can be regarded as spawning aggregations.

Co-occurrence of seagrass vegetation and coral colonies supports unique fish assemblages: a microhabitat-scale perspective.

Numerous studies have suggested that seagrass beds provide nursery habitats for juvenile fish in both tropical and subtropical regions. Most of these previous studies applied a landscape-scale perspective, in which seagrass beds and coral reefs are treated as independent, homogenous habitats. However, this perspective might overlook the microhabitat-scale perspective within the habitats, for example, the possibility that small-sized hard substrates (e.g., coral colonies) within seagrass beds might serve as fish nurseries. The present study aimed to examine the effects of the presence of microhabitats (small-sized coral colonies) within seagrass beds on the structure of fish assemblages. Fieldwork was conducted at Urasoko Bay, Ishigaki Island, Okinawa, Japan. Four habitat zones were selected: (1) seagrass bed with presence of massive coral colonies (SGCO), (2) seagrass bed without coral colonies (SG), (3) sandy bottom (without seagrass vegetation) with massive coral colonies (CO), and (4) sandy bottom without seagrass vegetation or coral colonies (SA). Six 20 m × 2 m line transects were established and monthly underwater observations were conducted between June and October in 2018 and 2019. A cluster analysis, analysis of similarity, and principal component analysis revealed that the fish assemblage at SGCO was significantly different from the other three habitat zones. This was because some fish species (e.g., Ostorhinchusishigakiensis and Lutjanus gibbus) were almost exclusively present at SGCO and rarely seen at CO, SG, and SA. Most individual fish belonging to these species were found on coral colonies at SGCO, suggesting that the co-occurrence of seagrass vegetation and coral colonies is essential for the habitats of these fish species. Although other fish species present at SGCO were also found at SG, three species, Parupeneus barberinus, Stethojulis strigiventer, and Lethrinus atkinsoni, were more abundant at SGCO with some found on coral colonies in this habitat zone. Several fish species that occurred at both SGCO and CO (e.g., Ostorhinchus properuptus, Cheilodispterus quinquelineatus, Chrysiptera cyanea, and Pomacentrus chrysurus) were more abundant or showed a greater size range at SGCO, suggesting greater survival rates in this habitat zone because of the co-occurrence of seagrass vegetation and coral colonies. This study demonstrated the existence of a unique fish assemblage structure at SGCO. Although the adoption of a landscape-scale perspective (three-dimensional structure of the vegetation) is necessary, a microhabitat-scale perspective that includes the presence of small hard substrates should also be considered to accurately evaluate the nursery function of seagrass beds.

Spatial distribution of parrotfishes and groupers in an Okinawan coral reef: size-related associations in relation to habitat characteristics.

Parrotfishes (Labridae: Scarini) and groupers (Epinephelidae) are important fish groups that are regarded as the fisheries targets of primary importance in coral reefs. In order to establish ecosystem-based management of these two fish groups, clarifying the spatial distribution relative to habitat characteristics is of central importance. The present study investigated the spatial distributions of 12 parrotfishes species and seven groupers species in relation to environmental characteristics in an Okinawan coral reef. Ten out of the 12 parrotfish species and all seven grouper species showed species-specific spatial distributions. Four substrate types in the inner reefs (branching Acropora, bottlebrush Acropora, dead branching Acropora, and dead bottlebrush Acropora), three substrate types in the exposed reefs (massive coral, other coral, and calcium carbonate substratum), and water depth showed significant associations with the spatial distribution of fishes. Among the 12 parrotfish species, two species (Scarus spinus and S. forsteni) and four species (S. psittacus, S. hypselopterus, S. dimidiatus and S. ghobban) were primarily found in exposed reefs and inner reefs, respectively. Among the seven grouper species, two species (Cephalopholis argus and C. urodeta) and two other species (C. miniata and Epinephelus ongus) were primarily found in exposed reefs and inner reefs, respectively. Size-related spatial distribution was also found for three parrotfish species (Chlorurus microrhinos, Scarus rivulatus and S. hypselopterus), indicating that smaller-sized and larger-sized individuals were respectively found at sites with greater coverage of substrates with fine structure (live bottlebrush Acropora and dead bottlebrush Acropora) and coarse structure (live branching Acropora, dead branching Acropora and calcium carbonate substratum). The present study suggested that the spatial distribution of parrotfishes and groupers is not necessarily associated with the higher coverage of living corals, but positively associated with high substrate complexity. Thus, actual spatial distributional patterns of species should be considered to select candidate sites for protection and conservation for the two fish groups.

Spatial distribution and feeding substrate of butterflyfishes (family Chaetodontidae) on an Okinawan coral reef.

Coral reefs support diverse communities, and relationships among organisms within these communities are quite complex. Among the relationships, clarifying the habitat association and foraging substrate selection relative to habitat characteristics is of central importance to ecology since these two aspects are the fundamentals for survival and growth of organisms. The aims of the present study were to investigate the spatial distribution and feeding substrate selection of 14 species of butterflyfishes on an Okinawan coral reef in Japan. Species-specific spatial distributions varied with habitat characteristics (e.g., encrusting corals, massive corals, branching Acropora and rock). For feeding substrates, seven species of obligate coral polyp feeders exhibited significant positive selectivity for tabular Acropora, corymbose Acropora, encrusting corals and massive corals but significant negative selectivity for dead corals, coral rubble and rock. Among six species of facultative coral polyp feeders, two species exhibited significant positive selectivity for encrusting corals and massive corals, and one species showed significant positive selectivity for dead corals as feeding substrates. In contrast, three species exhibited no significant positive selectivity for any feeding substrates. A similar result was observed for one non-coralline invertebrate feeder. Among the 14 species, 12 species showed a relatively close relationship between spatial distribution and feeding substrates but the remaining two species did not. The present study is the first study to elucidate species-specific spatial distributions and feeding substrate selection of butterflyfishes on an Okinawan coral reef. The results of the present study suggest that diverse substrates, including various types of living corals (especially encrusting corals, massive corals, tabular Acropora, corymbose Acropora and branching Acropora) and non-coralline substrates (rock) are the primary determinants of spatial distributions and feeding sites. Thus, diverse substrates are important for maintaining high species diversity of butterflyfishes and changes of the substrates would likely change the spatial patterns and foraging behavior, although species-specific responses may be found, depending on their species-specific dependence on vulnerable substrates.

Spatial distributions, feeding ecologies, and behavioral interactions of four rabbitfish species (Siganus unimaculatus, S. virgatus, S. corallinus, and S. puellus).

Clarifying the underlying mechanisms that enable closely related species to coexist in a particular environment is a fundamental aspect of ecology. Coral reefs support a high diversity of marine organisms, among which rabbitfishes (family Siganidae) are a major component The present study aimed to reveal the mechanism that allows rabbitfishes to coexist on coral reefs in Okinawa, Japan, by investigating the spatial distributions, feeding ecologies, and behavioral interactions of four species: Siganus unimaculatus, S. virgatus, S. corallinus, and S. puellus. All four species had a size-specific spatial distribution, whereby small individuals were found in sheltered areas that were covered by branching and bottlebrush Acropora spp. and large individuals were found in both sheltered and exposed rocky areas. However, no clear species-specific spatial distribution was observed. There was some variation in the food items taken, with S. unimaculatus primarily feeding on brown foliose algae, red foliose algae, and red styloid algae, and S. virgatus and S. puellus preferring brown foliose algae and sponges, respectively. However, S. corallinus did not show any clear differences in food preferences from S. virgatus or S. unimaculatus, mainly feeding on brown foliose algae and red styloid algae. The four species exhibited differences in foraging substrate use, which was probably related to differences in their body shape characteristics: S. unimaculatus has a slender body with a remarkably protruding snout and mainly used concave substrates for feeding, whereas S. virgatus has a deeper body with a low degree of snout protrusion and mainly used convex substrates. The other two species have a low degree of snout protrusion combined with a deeper body in the case of S. corallinus and a slender body in the case of S. puellus and used concave, flat, and convex substrates to an equal degree for feeding. Behavioral interactions were categorized into "agonistic behaviors" (attack and agonistic displays) and "no interactions." For all four species, a greater frequency of agonistic behaviors was observed when two conspecific pairs approached each other than when two heterospecific individuals encountered each other. Together, these results suggest that food item partitioning is one of the main factors enabling the coexistence of these four syntopic rabbitfish species, which is enhanced by species-specific differences in feeding substrates as a result of their different body shape and behavioral characteristics.

Spawning aggregation of white-streaked grouper Epinephelus ongus: spatial distribution and annual variation in the fish density within a spawning ground.

White-streaked grouper (Epinephelus ongus) is an important fisheries target and forms spawning aggregations at particular spawning grounds. The aims of the present study were to investigate the ecological characteristics of annual spawning aggregations such as (1) spatial variations in the density of E. ongus at the spawning ground, (2) the relationship between fish density and environmental variables, (3) inter-annual variations in the spawning aggregation, (4) the proportion of males to females at the spawning ground for several days pre-and post-spawning and (5) the relationship between male density and female density at the protected spawning ground, based on observations over five years at an Okinawan coral reef. Although the protected spawning ground area was large (ca. 2,500 m × 700 m), high density of E. ongus (over 25 individuals per 100 m2) was found in a limited area (within c.a. 750 m × 50 m). Current velocity and coverage of rocks had significant positive effects on the spatial distribution of E. ongus at the spawning ground. Inter-annual variation in the degree of aggregation was found and this variation was explained by the annual variation of mean seawater temperature during 40 days before the spawning day. The male-female ratio (male:female) at the spawning ground was ca. 3:1 for three years (May 2012, May 2014 and May 2015) whereas >13:1 for one year (May 2013). Significant positive relationships between male density and female density were found at the aggregation sites. It is suggested that E. ongus use aggregation sites with greater current velocity to reduce the risk of egg predation and seawater temperature is one of the main factors that is responsible for determining the degree of aggregation. It is also suggested that females possibly select sites with a greater density of males and this selection behavior might be the reason why females arrived at the spawning ground after the arrival of the males. For effective management of spawning grounds, precise site selection as well as the duration of the protection period are suggested to be key aspects to protect the spawning aggregations of E. ongus, which have been currently achieved at the spawning ground.

Parrotfish grazing ability: interspecific differences in relation to jaw-lever mechanics and relative weight of adductor mandibulae on an Okinawan coral reef.

Parrotfishes (family Labridae: Scarini) are regarded to have important roles for maintaining the ecosystem balance in coral reefs due to their removal of organic matter and calcic substrates by grazing. The purpose of the present study was to clarify the interspecific differences in grazing ability of five parrotfish species (Chlorurus sordidus, C. bowersi, Scarus rivulatus, S. niger and S. forsteni) in relation to interspecific differences in jaw-lever mechanics and the relative weight of the adductor mandibulae (muscles operating jaw closing). The grazing ability was calculated by using stomach contents (CaCO3 weight/organic matter weight) defined as the grazing ability index (GAI). There were significant interspecific differences in GAI (C. sordidus = C. bowersi > S. rivulatus > S. niger = S. forsteni). Teeth of C. sordidus and C. bowersi were protrusive-shape whereas teeth of S. rivulatus, S. niger and S. forsteni were flat-shape. C. sordidus and C. bowersihave jaw-lever mechanics producing a greater biting force and have a larger weight of adductor mandibulae. S. rivulatus has jaw-lever mechanics producing a greater biting force but a smaller weight of adductor mandibulae that produce an intermediate biting force. In contrast, S. niger and S. forsteni have jaw-lever mechanics producing a lesser biting force and have a smaller weight of adductor mandibulae. Feeding rates and foray size of S. rivulatus, S. niger and S. forsteni were greater than C. sordidus and C. bowersi. The degree in bioerosion (GAI × feeding rate) was the largest for S. rivulatusand the smallest for S. forsteni. The degree in bioerosion for C. sordidus was larger than S. niger whereas relatively equal between C. bowersi and S. niger. These results suggest that interspecific difference in GAI was explained by interspecific differences in teeth shape, jaw-lever mechanics and relative weight of adductor mandibulae. The interspecific difference in the degree of bioerosion suggests the importance of various size of parrotfishes with diverse feeding modes to maintain healthy coral reef ecosystems.

Pair formation, home range, and spatial variation in density, size and social status in blotched foxface Siganus unimaculatus on an Okinawan coral reef.

The present study examined pair formation, spatial pattern of home range and spatial variation in density, size and social status of blotched foxface Siganus unimaculatus (family Siganidae) on an Okinawan coral reef. Of 32 pairs sampled for sexing, 31 (96.9%) were heterosexual and showed size-assortative pairing. Developed ovaries were found in April and July, whereas oocytes were immature in August, September and February. Heterosexual pairing was found in both reproductive and non-reproductive periods. Home range size tended to be positively related to fork length (FL). The degree of home range overlap for same size class pairs was smaller than that for different size class pairs. The intraspecific behavior when two pairs approached each other was categorized as 'attack,' 'agonistic display' and 'no interactions,' and the frequency of agonistic behaviors ("attack" or "agonistic display") was significantly greater than "no interactions." Underwater observations at a seagrass bed, a rocky reef flat and a sheltered reef slope revealed that the mean FL was significantly smaller at the sheltered reef slope (4-13 cm) than at the rocky reef flat (>13 cm). No individuals were found in the seagrass bed. Most individuals less than 6 cm FL were solitary, whereas most individuals over 7 cm FL were paired. Density was significantly greater on the sheltered reef slope than on the rocky reef flat.

Use of a gyroscope/accelerometer data logger to identify alternative feeding behaviours in fish.

We examined whether we could identify the feeding behaviours of the trophic generalist fish Epinephelus ongus on different prey types (crabs and fish) using a data logger that incorporated a three-axis gyroscope and a three-axis accelerometer. Feeding behaviours and other burst behaviours, including escape responses, intraspecific interactions and routine movements, were recorded from six E. ongus individuals using data loggers sampling at 200 Hz, and were validated by simultaneously recorded video images. For each data-logger record, we extracted 5 s of data when any of the three-axis accelerations exceeded absolute 2.0 g, to capture all feeding behaviours and other burst behaviours. Each feeding behaviour was then identified using a combination of parameters that were derived from the extracted data. Using decision trees with the parameters, high true identification rates (87.5% for both feeding behaviours) with low false identification rates (5% for crab-eating and 6.3% for fish-eating) were achieved for both feeding behaviours.

On the fourth Diadema species (Diadema-sp) from Japan.

Four long-spined sea urchin species in the genus Diadema are known to occur around the Japanese Archipelago. Three species (D. savignyi, D. setosum, and D. paucispinum) are widely distributed in the Indo-Pacific Ocean. The fourth species was detected by DNA analysis among samples originally collected as D. savignyi or D. setosum in Japan and the Marshall Islands and tentatively designated as Diadema-sp, remaining an undescribed species. We analyzed nucleotide sequences of the cytochrome oxidase I (COI) gene in the "D. savignyi-like" samples, and found all 17 individuals collected in the mainland of Japan (Sagami Bay and Kyushu) to be Diadema-sp, but all nine in the Ryukyu Archipelago (Okinawa and Ishigaki Islands) to be D. savignyi, with large nucleotide sequence difference between them (11.0%±1.7 SE). Diadema-sp and D. savignyi shared Y-shaped blue lines of iridophores along the interambulacrals, but individuals of Diadema-sp typically exhibited a conspicuous white streak at the fork of the Y-shaped blue iridophore lines, while this feature was absent in D. savignyi. Also, the central axis of the Y-shaped blue lines of iridophores was approximately twice as long as the V-component in D. savignyi whereas it was of similar length in Diadema-sp. Two parallel lines were observed to constitute the central axis of the Y-shaped blue lines in both species, but these were considerably narrower in Diadema-sp. Despite marked morphological and genetic differences, it appears that Diadema-sp has been mis-identified as D. savignyi for more than half a century.