Diurnal periodicity of spawning activity of permanently territorial damselfishes (Teleostei: Pomacentridae)

Synopsis Like other congeneric damselfishes, the herbivorous Stegastes altus defends individual feeding territories from heterospecific food competitors, regardless of sex. Females spawned demersal eggs for 31.4 min (n = 25) at the nest in territories of males 0–75 m away from theirs. Throughout the spawning season (June to September), spawning occurred only at dawn, mainly just after sunrise. Daily activity of the fish community showed that potential diurnal food competitors were few or inactive only at dawn and dusk. The frequency of intrusions into the female's territory by heterospecific competitors were as low when she left her territory to spawn at dawn, as they were in the daytime when she defended it. Removal of the female in the daytime resulted in a significant increase in intrusion frequency. An ‘anti-competitor hypothesis’, whereby dawn spawning ensures the food resource in the female's territory seems to explain not only the spawning periodicity in S. altus but also the timing of spawning of other permanently territorial damselfishes. Contribution from the Laboratory of Animal Ecology, Kyoto University, No. 492.     

Evolution and diversity of ram-suction feeding in damselfishes (Pomacentridae)

The cerato-mandibular (c-md) ligament is a synapomorphy within Pomacentridae that creates a tight link between the lower jaws and the hyoid bars. However, this morphological trait has been secondarily lost in multiple lineages during evolution. A previous study revealed that the loss of this trait acted as a release of evolutionary constraints, leading to a cascade of morphological changes such elongated buccal jaws and a slender body. Ecomorphological interpretations suggested the loss of the c-md ligament has ultimately led to a new adaptive peak in zooplanktivory through an optimization of the ram feeding mode associated with a specialization in pelagic feeding. Here, we tested these hypotheses by comparing functional and diet diversity between damselfish species with and without the c-md ligament. Although species lacking the c-md ligament presented a conserved kinematic pattern resulting from high ram and low suction performances, our results did not support an optimization of the ram feeding mode. Indeed, some species with the c-md ligament showed the same or exceeded the ram performance of species without the c-md ligament. The species with the c-md ligament had a more diverse kinematic pattern exploring the entire ram-suction functional range in damselfishes. Finally, our results did not show any diet variations associated with the loss of the c-md ligament. Our study furthers the understanding of how a morphological trait has shaped, by its presence or absence, the ecomorpho-functional diversification of Pomacentridae.     

Comparative trophic morphology in eight species of damselfishes (Pomacentridae)

Damselfishes show significant biodiversity in the coral reefs. To better understand such diversity, an ecomorphological approach was investigated in the trophic morphology of eight species of Pomacentridae (Chromis acares, C. margaritifer, Dascyllus aruanus, D. flavicaudus, Pomacentrus pavo, Plectroglyphidodon johnstonianus, Pl. lacrymatus and Stegastes nigricans) belonging to different trophic guilds (zooplankton, algal, coral polyp feeders and omnivores). Geometric morphometrics were used to quantify size and shape variations in four skeletal units: (1) neurocranium, (2) suspensorium and opercle, (3) mandible and (4) premaxilla. This method allowed us to reveal shape and size differences correlated to functional diversity both within and between trophic guilds. Among zooplanktivores, C. margaritifer, D. aruanus and D. flavicaudus have a high and long supraoccipital crest, short mandibles forming a small mouth and high suspensoria and opercles. These three species can be considered to be suction feeders. In the same guild, C. acares shows opposite characteristics (long and thin mandibles, lengthened neurocranium and suspensorium) and can be considered as a ram feeder. Among herbivores and corallivores, the two species of Plectroglyphidodon and S. nigricans can be considered as grazers. Differences in skeletal shape are mainly related to improving the robustness of some skeletal parts (broad hyomandibular, short and high mandibles). The shapes of P. pavo, which feeds largely on algae, strongly differ from that of the other three grazers exhibiting similar morphological characteristics to C. acares (e.g., long and shallow suspensorium, lengthened neurocranium). This highlights likely differences concerning cutting or scraping method. Finally, no strong correlations exist between size and shapes in the eight studied species. Size difference among species having a very similar shape could be viewed as a factor optimizing resource partitioning.     

Echinoderms (Echinodermata) from the Gulf of California, Mexico

A systematic list of the echinoderms of the Gulf of California, based on museum specimens of the Colección Nacional de Equinodermos, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autónoma de México and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. is presented. A total of 193 echinoderm species is recorded, distributed in 108 genera, 51 families and 19 orders. 12 new records for the Gulf of California are presented: Asteroidea (four), Ophiuroidea (three) and Holothuroidea (five).     

Bergmann ecogeographic trends among triplefin blennies (Teleostei: Tripterygiidae) in the Gulf of California,Mexico

Synopsis Bergmann-type ecogeographic patterns are common among small resident rocky-shore fishes in the Gulf of California where triplefin blennies (Tripterygiidae) represent a significant component of the community. Using estimates of adult body size from samples throughout the Gulf, four triplefin species, three more than previously reported, showed significant correlations of body size with latitude. The intensity and form of the relationships are given. Measuring the length of the largest individual in samples rather than using average body weight per individuals in samples offers a better estimate of adult body size.     

Habitat use and behavioural ecology of the juveniles of two sympatric damselfishes (Actinopterygii: Pomacentridae) in the south-western Atlantic Ocean

Spatial distribution, microhabitat use and territorial and feeding behaviours were compared between the juveniles of two sympatric territorial damselfishes Stegastes variabilis and Stegastes fuscus on a small tropical reef in the south-western Atlantic Ocean. Juvenile S. variabilis were most abundant at sites subject to stronger hydrodynamics and with mixed benthic cover, whereas juvenile S. fuscus were most abundant at sheltered sites with dense turf algae cover. No differences regarding feeding habits were detected, with both species preferentially feeding on turf algae. Also, despite similarities in territory area and agonistic encounter rates, the identity and proportion of intruders involved in agonistic interactions differed significantly between species. These interdependent traits suggest strong asymmetric competition, with juvenile S. fuscus dominating high-quality sites and evicting juvenile S. variabilis to low quality, marginal areas of the reef.     

Molecular systematics of the damselfishes (Teleostei: Pomacentridae): Bayesian phylogenetic analyses of mitochondrial and nuclear DNA sequences

Damselfishes in the family Pomacentridae represent one of the few families of reef fishes found on coral reefs irrespective of location. At a local scale, damselfishes are often the most abundant coral reef fish, and their study has provided much of our current understanding of the ecology of tropical reef animals. The study of phylogenetic relationships among the Pomacentridae has lagged ecological investigation of the group, thus limiting historical perspective on the remarkable species richness of the family. In this study, we used 1989bp of DNA sequence representing three mitochondrial genes and 1500bp of the single copy nuclear RAG1 region to infer hypotheses of relationship for the group. Our analysis includes 103 Pomacentridae species in 18 genera, and three of the four named subfamilies: Amphriprioninae, Chrominae, and Pomacentrinae. The Bayesian method of phylogenetic reconstruction was applied to the data, because even with a large number of sequences it is an efficient means of analysis that provides intuitive measures of support for tree topologies and for the parameters of the nucleotide substitution model. Four Pomacentridae clades were identified with high statistical support whether the data were analyzed from a mtDNA, RAG1 or combined perspective, and in all analyses the current subfamilial classification of the Pomacentridae was rejected. At the genus level, Amphiprion, Chromis, and Chrysiptera were also rejected as natural groups. Abudefduf, Amblyglyphidodon, Dascyllus, Neoglyphidodon, Neopomacentrus, and Pomacentrus were each strongly supported as monophyletic genera but the support for monophyly is nonetheless compromised by sample size, except in the case of Dascyllus and Abudefduf for which we have sampled almost all of the described species.     

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.     

Reproductive activity and condition index of Holacanthus passer (Teleostei: Pomacanthidae) in the Gulf of California,Mexico

Annual variation and relationship between the gonadosomatic index (IG) and the condition index (K) of Holacanthus passer were evaluated in 194 specimens from Cueva de León, Gulf of California (24 degrees 02' N, 110 degrees 24' W). There was a significant negative correlation between IG and K in both sexes: indices exhibit an opposite seasonal pattern. The reproductive activity of both sexes is inversely related with nutritional status. There was a significant difference in IG but not in K values between males and females. The condition index may be used to determine reproduction time of the species in other localities without sacrificing the organisms. This could be a valuable tool to develop monitoring programs for fisheries and culture of this species.     

Deep-water polychaetes (Annelida) from the southeastern Gulf of California, Mexico

Polychaetes inhabiting deep-sea soft bottoms from the southeastern Gulf of California were collected during four oceanographic cruises during 2000 and 2001. Sampling of benthic organisms was performed with a benthic sledge to collect epifauna and a Karling dredge for epifauna and infauna, in a depth range from 732 to 2 250 m. A list of the polychaetes that were collected and their distribution are presented here. A total of 73 species (distributed among 33 families) were identified. Moreover, 11 species were identified only to genus level and 20 species only to family level. With the exception of Ancistrosyllis hartmanae and Melinnampharete eoa, all identified species have been previously reported in soft bottoms of the Gulf of California or in adjacent areas. Additional previously unreported information is provided herein regarding depth ranges, geographical distribution, morphology and tubes inhabited by the organisms. The morphology of the ampharetids Amage sp. and Samytha sp. does not coincide with that of other species in these genera reported for the Gulf of California, which suggests that they are probably undescribed species.     

Modular diversification of the locomotor system in damselfishes (Pomacentridae)

As fish move and interact with their aquatic environment by swimming, small morphological variations of the locomotor system can have profound implications on fitness. Damselfishes (Pomacentridae) have inhabited coral reef ecosystems for more than 50 million years. As such, habitat preferences and behavior could significantly constrain the morphology and evolvability of the locomotor system. To test this hypothesis, we used phylogenetic comparative methods on morphometric, ecological and behavioral data. While body elongation represented the primary source of variation in the locomotor system of damselfishes, results also showed a diverse suite of morphological combinations between extreme morphologies. Results show clear associations between behavior, habitat preferences, and morphology, suggesting ecological constraints on shape diversification of the locomotor system. In addition, results indicate that the three modules of the locomotor system are weakly correlated, resulting in versatile and independent characters. These results suggest that Pomacentridae is shape may result from the interaction between (1) integrated parts of morphological variation that maintain overall swimming ability and (2) relatively independent parts of the morphology that facilitate adaptation and diversification. J. Morphol. 277:603–614, 2016. © 2016 Wiley Periodicals, Inc.     

Agar properties of two species of Gracilariaceae from the Gulf of California,Mexico

Agar properties of two potentially commercial important seaweeds from the Gulf of California were studied. Maximum yield in Gracilaria vermiculophylla (45.7%) occurred during the summer months, coinciding with high water temperatures (31°C) whereas minimum yields (11.6%) were obtained during the coldest months of the year when populations of this species diminish in the bay. Gracilariopsis longissima showed two yield peaks, one in spring and another in fall, before the maximum and minimum seawater temperatures. Gel strength in native agar from the two species was low (<22.5 g cm⁻²) for most of the year. G. vermiculophylla native agar showed a slight increase in gel strength from June to August, which were the hottest months. Maximum value was 85 g cm⁻¹ in August. Maximum gel strength in G. longissima was observed in October (91 g cm⁻¹), and an unusual native agar with no detectable gel strength was observed in March and April samples. Gelling temperatures range from 27.7 to 36.5°C in G. vermiculophyla and from 26.6 to 34.9°C in G. longissima, meanwhile melting points were 73.9 – 53.5°C and 75.5 – 56.6°C, respectively. Sulfate content was high, 6.3–13.9% in G. vermiculophylla and 1.9–11.9% in G. longissima, and on the other hand 3,6 anhydrogalactose content was low 12.1–26.7% and 9.1–23%, respectively compared to other species. Results obtained showed that mean native agar yields of Gracilaria vermiculophylla and Gracilariopsis longissima from the Gulf of California are comparable to other tropical Gracilaria. However, the low gel strength, high sulfate content and low 3,6 anhydrogalactose content observed in the native agar extracted from these species make this an agaroid, thus alternative methods of extraction should be used to evaluate the possibility of commercial utilization of both species.     

Sound production in two species of damselfishes (Pomacentridae): Plectroglyphidodon lacrymatus and Dascyllus aruanus

Agonistic sounds of two pomacentrid species, Plectroglyphidodon lacrymatus and Dascyllus aruanus , were recorded in captivity. Plectroglyphidodon lacrymatus produced trains of 2–5 pops, each composed of 18–25 cycles, with an average duration of 56 ms; most energy ranged from c. 100 to 1000 Hz. Dascyllus aruanus produced pops and chirps. Pops were generally composed of a single pulse, with 2–14 peaks and an average duration of 6·7 ms. Pops contained energy >4 kHz, and peak frequency ranged from 680 to 1300 Hz. Chirps consisted of trains of 12–42 short pulses of three to six cycles, with durations varying from 0·6 to 1·27 ms; peak frequency varied from 3400 to 4100 Hz. Sound production in P. lacrymatus suggested that pomacentrids are derived from an ancestral taxon capable of sound production and that this capacity is a synapomorphy for the family. Although in the Pomacentridae, pops are typically composed of a single pulse, which is longer and higher pitched than chirps composed of a series of shorter pulses, D. aruanus chirps were higher pitched than its pops. Thus, acoustic variation in the genus Dascyllus is probably not more restricted than in the Pomacentridae.     

Behavior and physiology of coexistence in two species of Dascyllus (Pomacentridae,Teleostei)

Synopsis In the Gulf of Aqaba Dascyllus aruanus and D. marginatus, form distinct populations organised into harems of single males and groups of females, living permanently in Stylophora and Acropora corals. Each harem consists of 4 sexual casts: dominating males, adult females, sex-changing fish and juveniles. The sexual performance of each adult fish in such social units is controlled by the presence or absence of a dominating male. In corals occupied by mixed groups of both species, only those females spawn that belong to the same species as the dominating male. Direct observation in the sea shows that such situations are produced by indiscriminative settling mechanisms of groups of juvenile fish. Evidence is also presented, demonstrating that dispersion of coral colonies suitable for occupancy and predation by piscivorous fish can modulate such complicated reproduction.     

Comparative ecology of subtidal actiniarians from the coasts of California and the Gulf of Maine,USA

Shallow subtidal populations of sea anemones occupying hard substrates along the California (Pacific) and Gulf of Maine (Atlantic) coasts of the USA are compared. The diversity of anemones in the southern Gulf of Maine to 35 m is low (4 species) in contrast to similar habitats in California (at least 11 species), but parallel patterns of individual size, distribution and predation impacts occur. Water motion influences depth distribution and size, while predation affects size, population structure and habitat selection. The nudibranch Aeolidia papillosa exerts a similar pressure on population structure in some anemone species in both areas as a size selective predator. The seastar Dermasterias imbricata appears to magnify the selective pressure for large size in the eastern Pacific coastal zone. Interspecific aggression of anemones determines some distribution patterns in areas of overlap.     

Diversification of the pectoral fin shape in damselfishes (Perciformes, Pomacentridae) of the Eastern Pacific

Fin shape strongly influences performance of locomotion across all swimming styles. In this study, we focused on the diversity of the pectoral fin morphology in damselfishes of the Eastern Pacific. Underwater observations and a review of literature allowed the characterization of ten behavioral groups. Territorial and non-territorial species were discriminated easily with traditional morphometrics. Five ecomorphological groups were recognized by geometric morphometric analyses. Geometric data segregated the outgroup from the damselfishes and allowed the distinction of mean morphologies from extreme ones within territorial and non-territorial species. Additionally, geometric morphometric data split Abudefduf into two groups: (1) A. troschelii is similar to C. atrilobata and (2) A. concolor and A. declivifrons are close to Stegastes. Solitary territorial species (e.g., Stegastes) show rounded and high fins whereas non-territorial species living in groups (e.g., Chromis) present long and curved pectoral fins. In the range of morphological variation, the morphologies of Microspathodon (elongate with highly curved hydrodynamic trailing edge) and Azurina (long, slender and angular) represent the extreme morphologies within territorial and non-territorial species, respectively. Our study revealed a strong relationship between the pectoral fin shape and the behavioral diversification in damselfishes.     

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.