Ecological speciation in tropical reef fishes

The high biodiversity in tropical seas provides a long-standing challenge to allopatric speciation models. Physical barriers are few in the ocean and larval dispersal is often extensive, a combination that should reduce opportunities for speciation. Yet coral reefs are among the most species-rich habitats in the world, indicating evolutionary processes beyond conventional allopatry. In a survey of mtDNA sequences of five congeneric west Atlantic reef fishes (wrasses, genus Halichoeres) with similar dispersal potential, we observed phylogeographical patterns that contradict expectations of geographical isolation, and instead indicate a role for ecological speciation. In Halichoeres bivittatus and the species pair Halichoeres radiatus/brasiliensis, we observed strong partitions (3.4% and 2.3% divergence, respectively) between adjacent and ecologically distinct habitats, but high genetic connectivity between similar habitats separated by thousands of kilometres. This habitat partitioning is maintained even at a local scale where H. bivittatus lineages are segregated between cold- and warm-water habitats in both Bermuda and Florida. The concordance of evolutionary partitions with habitat types, rather than conventional biogeographical barriers, indicates parapatric ecological speciation, in which adaptation to alternative environmental conditions in adjacent locations overwhelms the homogenizing effect of dispersal. This mechanism can explain the long-standing enigma of high biodiversity in coral reef faunas.     

Cleaning interactions at the only atoll in the South Atlantic

In marine ecosystems, cleaning is a mutualistic relationship in which so-called cleaners remove ectoparasites, diseased tissue, or mucus from the body of their clients, and thus help to maintain a healthy reef community. In spite of its importance in many marine habitats, this interaction remains poorly understood, particularly at oceanic islands. Here, we present the first comprehensive study of cleaning interactions in a reef fish assemblage at Rocas, the only atoll in the South Atlantic. We recorded 318 cleaning events, in which six fish species, including two endemic ones, and two shrimp species acted as cleaners. The clients serviced by these cleaners were 21 bony fish species, one shark and one sea turtle. The cleaner wrasse Thalassoma noronhanum and the cleaner goby Elacatinus phthirophagus were the cleaners with the greatest number of events and species richness of clients. Additionally, 82% of clients in the cleaning events were non-piscivores, and the abundance of both cleaners and clients positively influenced the number of cleaning events (R² = 0.4; p < 0.001). Our results indicate that Rocas atoll has a high species richness of cleaner species despite its small size and highlight the importance of studies of cleaning symbiosis, even in isolated places with low species richness, for a better comprehension of this association in reefs.     

Diel feeding migrations in tropical reef fishes

Summary 1. Many tropical reef fishes that feed during the day rest at night, whereas many that feed at night rest during the day. The feeding grounds of many are some distance from their resting grounds. Thus they migrate between these two locations during twilight as part of a general changeover between diurnal and nocturnal situations.2. At least many of these migrations are predictable, both as to time and to the route taken. The distances traveled vary between species, ranging from just a few meters, to more than several kilometers.3. The pattern of migrations is strongly influenced by the relative threat from predators at different periods of the diel cycle.4. During the day, migrations of reef fishes are limited to intra-reef movements: short vertical movements by certain plankton feeders, and lateral excursions from one part of the reef to another by certain herbivores and plankton feeders. Movements into the open regions that lie adjacent to many reefs are not adaptive in daylight due largely to a danger from predators.5. Despite constant threat from predators during the day, smaller reef fishes remain relatively secure during most of this period by staying close to shelter, or by schooling. However, these defenses are less effective during twilight, when the danger from predators intensifies. The diurnal migrators return to the shelter of their resting places prior to that part of evening twilight when danger is greatest, and the nocturnal migrators usually do not expose themselves for their nightly foraging until after the period of maximum danger has passed. During morning twilight the sequence is reversed.6. The major mechanisms whereby smaller reef fishes reduce predation during the day — schooling and staying close to shelter — are less evident at night. Not only do reef fishes range freely at night into the open regions that are avoided in daylight, but their schools are more loosely defined, and many are active as solitary individuals or in small groups. The tendency for looser associations and ranging farther afield increases on darker nights.7. Most predators that threaten reef fishes are visual feeders whose mode of attack loses effectiveness when light falls below a certain level. Although they operate to some extent under moonlight, they threaten small reef fishes less at night than during the day.8. In addition to whatever other ways a shcool may be adaptive, by reducing variable behavior among its members the school is especially important to migrating species. Responses to the various cues that mark the migration routes may be refined to within acceptable limits for the population as a whole only by coordinated group action.9. Submarine topographical features are important reference points for migrating reef fishes.

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Tägliche Futtersuchwanderungen bei tropischen Riffischen

Kurzfassung Dämmerungszeiten wandern viele tropische Fische zwischen ihren Ruheplätzen auf dem Riff und ihren Futterplätzen. Diese Wanderungen, die sowohl von tag- als auch von nachtaktiven Fischen durchgeführt werden, sind bedingt durch den Licht-Dunkel-Wechsel, und für viele Arten können sogar Zeit und Weg der Wanderung vorhergesagt werden. Das Muster der tagesrhythmischen Wanderungsbewegungen wird vor allem durch das Ausmaß von Bedrohungen durch Raubfeinde beeinflußt. Bewegungen in ungeschützte Gebiete, die vom Riff wegführen, sind am Tage nicht vorteilhaft. Bei Einbruch der Dunkelheit sind diese offenen Gebiete hingegen ein Konzentrationspunkt der Wanderungen aus dem Riff, wobei sich auch der Zusammenhalt der Fischschwärme verringert. Da Schwarmverhalten und Schutzsuchen die beiden wichtigsten Verteidigungseinrichtungen gegenüber Räubern darstellen, bedeutet das Aufgeben dieser Verhaltensweisen zu Beginn der Dämmerung, daß während der Nacht kleinere Riffische weniger bedroht sind als während des Tages. Für wandernde Schwarmfische dürfte eine geringe Variabilität der Verhaltensweisen von adaptiver Bedeutung sein. Nur durch ein koordiniertes Schwarmverhalten ist es offensichtlich möglich, innerhalb bestimmter Grenzen auf die verschieden optischen Marken, welche die Wanderwege kennzeichnen, zu reagieren. Vermutlich spielen die topographischen Gegebenheiten unter Wasser eine entscheidende Rolle für die Orientierung der Riffische.

     

A comparative study of cleaning activity of two reef fishes at Fernando de Noronha Archipelago,tropical West Atlantic

Cleaner fishes are usually classified as obligate or facultative cleaners according to their diet and the extent to which their nutritional requirements in the different ontogenetic stages are gained from cleaning. While obligate cleaners clean throughout their lives and ingest mainly food taken from the clients’ body surface, facultative cleaners clean only as juveniles and have a broader diet. In addition, some facultative cleaners may experience a relatively higher predation risk, and thus rarely interact with piscivorous fishes. Despite these acknowledged differences, there are very few studies that compare cleaning activity of obligate and facultative cleaners within the same area. Cleaning activity of the obligate cleaner goby Elacatinus cf. randalli and the facultative cleaner wrasse Thalassoma noronhanum were comparatively examined at Fernando de Noronha Archipelago, tropical West Atlantic. The client assemblage attended by the two cleaners differed, as the goby attended a slightly greater diversity of species (22), mostly piscivores and zoobenthivores, and the wrasse attended fewer species (19), mostly planktivores. Chromis multilineata was the most common client species of both cleaners, although body size (which is expected to be positively correlated to clients’ ectoparasite load) of C. multilineata individuals attended by the goby was larger than that of the individuals attended by the wrasse. Despite such differences, T. noronhanum showed a surprisingly species-rich client assemblage when compared with other cleaners of the genus Thalassoma. In addition, the frequency and time spent on cleaning interactions, as well as the number of client species attended per 10-min period, was similar for both cleaner species, which indicate that they have important yet complimentary ecological roles in the reef community at Fernando de Noronha Archipelago.     

Non-random reef use by fishes at two dominant zones in a tropical,algal-dominated coastal reef

Habitat use and the processes which determine fish distribution were evaluated at the reef flat and reef crest zones of a tropical, algal-dominated reef. Our comparisons indicated significant differences in the majority of the evaluated environmental characteristics between zones. Also, significant differences in the abundances of twelve, from thirteen analyzed species, were observed within and between-sites. According to null models, non-random patterns of species co-occurrences were significant, suggesting that fish guilds in both zones were non-randomly structured. Unexpectedly, structural complexity negatively affected overall species richness, but had a major positive influence on highly site-attached species such as a damselfish. Depth and substrate composition, particularly macroalgae cover, were positive determinants for the fish assemblage structure in the studied reef, prevailing over factors such as structural complexity and live coral cover. Our results are conflicting with other studies carried out in coral-dominated reefs of the Caribbean and Pacific, therefore supporting the idea that the factors which may potentially influence reef fish composition are highly site-dependent and variable.     

Ecological traits influencing range expansion across large oceanic dispersal barriers: insights from tropical Atlantic reef fishes

How do biogeographically different provinces arise in response to oceanic barriers to dispersal? Here, we analyse how traits related to the pelagic dispersal and adult biology of 985 tropical reef fish species correlate with their establishing populations on both sides of two Atlantic marine barriers: the Mid-Atlantic Barrier (MAB) and the Amazon–Orinoco Plume (AOP). Generalized linear mixed-effects models indicate that predictors for successful barrier crossing are the ability to raft with flotsam for the deep-water MAB, non-reef habitat usage for the freshwater and sediment-rich AOP, and large adult-size and large latitudinal-range for both barriers. Variation in larval-development mode, often thought to be broadly related to larval-dispersal potential, is not a significant predictor in either case. Many more species of greater taxonomic diversity cross the AOP than the MAB. Rafters readily cross both barriers but represent a much smaller proportion of AOP crossers than MAB crossers. Successful establishment after crossing both barriers may be facilitated by broad environmental tolerance associated with large body size and wide latitudinal-range. These results highlight the need to look beyond larval-dispersal potential and assess adult-biology traits when assessing determinants of successful movements across marine barriers.     

New trematode genera from fishes of the southern Atlantic

Two new genera of the lepocreadiid trematodes, Gibsonia gen. n., (type species--G. hastata sp. n.) from Macrourus carinatus Günther, 1878 (Macrouridae), collected in the Falkland Islands area, and Muraenolepitrema gen. n. (type species--M. magnatestis sp. n.) from Muraenolepis microps L?nnberg, 1905 (Muraenolepidae), collected in the South Georgia Island area, are described. Gibsonia gen. n. differs from all known genera of the family Lepocreadiidae by unusual body shape divided into 2 parts, suctorial formations, great number of glandular cells and a muscular bulge of the body sides; it differs from the closely related genus Lepidapedon by separate genital pores. Muraenolepitrema gen. n. differs from Myzoxenus by bursa shape, separate genital pores, lobed shape of genital organs.     

Two new anthiine fishes from the eastern tropical Atlantic

Two anthiine fishes from the eastern tropical Atlantic are described as newHolanthias cyprinoides andAnthias helenensis. The former is distinguished from the other Atlantic species ofHolanthias in having the forked caudal fin with rounded lobes and from the Indo-Pacific species in having no elongated dorsal spines or soft rays. The latter is closely related toAnthias asperilinguis Günther (South America, Atlantic coast), but differs from it in having more pectoral fin rays and more gill rakers. The present investigation onAnthias suggests that AtlanticAnthias is a genus distinct fromPseudanthias of the Indo-Pacific.     

Ecomorphological correlates in tropical stream fishes of southern India

Synopsis I studied ecomorphological correlations in 18 stream fish species (belonging to five families) commonly occurring in diversity rich habitats of rivers in the central Western Ghats (Southern India). Univariate and multivariate analyses indicated a clear segregation of species in morphological and ecological (habitat) space. Cluster analyses using morphological features segregated species based mainly on their body shape and feeding parameters while habitat parameters segregated species according to their position along the water column and preference to runs or pool habitats. Principal component analyses showed that body size, head dimensions, mouth position and eye diameter were important characters for species segregation in morphological space while run or pool habitat type and the position of the fish in the water column best explained segregation of species in habitat space. Consistencies in some of the species clusters obtained from analyses on morphological characteristics and habitat preferences warrant closer examination of the relationships between them. These were studied employing multivariate methods like canonical correspondence analysis and the Mantel test. The tests revealed weak correlations indicating that in addition to morphology, other factors like behavioural, physiological and evolutionary constraints together are likely to play an important role in ecosystem structuring of these complex tropical assemblages.     

Theory of resource allocation strategy in territorial male tropical reef fishes

A mathematical model is proposed to explain energy resource allocation between sperm production and territoriality in male reef fishes (Labridae species) from the point of view of optimization. Labridae species are typically characterized by both TP (terminal phase) and IP (initial phase) males. The former are considered to release a lower amount of sperm but show aggressive territoriality. In the model, TP male reproductive success is considered as depending upon both fertilization probability (depending on sperm density) and the individual's own territorial activities. Between these factors, a trade-off exists by which the fertilization probability can be enhanced only by reducing territoriality. Therefore, the male has to decide how much of the total available energy resource should be allocated to each. The model showed that under high fertilization efficiency the male can achieve high success by spending less of the resource on sperm production and correspondingly more for territoriality. The TP male reproductive success increases with decreasing male density in the habitat. Nevertheless, when intruding males cannot be excluded completely by territorial behavior of the TP male, females prefer high male density. If females can control the number of intruding males to some degree, conflict may arise between the sexes. Received: December 8, 1999 / Accepted: May 24, 2000     

Habitat use patterns of fishes across the mangrove-seagrass-coral reef seascape at Ishigaki Island,southern Japan

To clarify seascape-scale habitat use patterns of fishes in the Ryukyu Islands (southern Japan), visual censuses were conducted in the mangrove estuary, sand area, seagrass bed, coral rubble area, branching coral area on the reef flat, and tabular coral area on the outer reef slope at Ishigaki Island in August and November 2004, and May, August and November 2005. During the study period a total of 319 species were observed. Species richness and abundance were highest in the branching and tabular coral areas, followed in order by the seagrass bed and mangrove estuary, and coral rubble and sand areas, in each month. Cluster analysis resulted in a clear grouping of assemblage structures by habitat type rather than by census month. SIMPER analysis showed that fish assemblages in the tabular coral area were mainly characterized by Acanthurus nigrofuscus, Pomacentrus lepidogenys, P. philippinus and P. vaiuli, the branching coral area by Chromis viridis and Pomacentrus moluccensis, the coral rubble area by Amblyeleotris steinitzi and Ctenogobiops pomastictus, the seagrass bed by Cheilio inermis, Lethrinus atkinsoni and Stethojulis strigiventer, the sand area by Valenciennea longipinnis, and the mangrove estuary by Gerres oyena, Lutjanus fulvus and Yongeichthys criniger. Moreover, fishes exhibited two habitat use strategies, inhabiting either a single or several specific habitats throughout their benthic life history stages, or having a possible ontogenetic habitat shift from the mangrove estuary or seagrass bed to coral-dominated habitats (e.g., Lethrinus atkinsoni, Lethrinus obsoletus, Lutjanus fulviflamma, Lutjanus fulvus, Lutjanus gibbus, Lutjanus monostigma and Parupeneus barberinus), suggesting that the mangrove estuary and seagrass bed have a nursery function.     

Egg size in relation to fertilization dynamics in free-spawning tropical reef fishes

In marine invertebrates that spawn by simply releasing their gametes into the water (free-spawning), fertilization success likely is often limited by low sperm concentrations, due to dispersion of mates and dispersal of gametes by water movements. Production of large, low density eggs might be advantageous when sperm concentrations consistently are low, because large target size might increase egg/sperm encounters, and more low than high density eggs could be produced per clutch. Although average fertilization success in the labrid Thalassoma bifasciatum is 95% in both group spawns (in which multiple males compete for fertilizations by producing large quantities of sperm) and pair (mono-male) spawns, it is slightly lower in pair spawns, due to low level sperm limitation that arises because pair-spawning males release near the minimum number of sperm necessary for maximum fertilization. I examined whether variation in egg size and content in T. bifasciatum and other free-spawning fishes is related to variation in spawning mode, to assess whether compensatory production of large, low-density eggs might be contributing to high fertilization success in pair spawns. I found no difference between the volume or density of eggs of (1) pair- and group-spawning females of T. bifasciatum, or (2) pair-and group-spawning congeneric species of labrids, scarids, and serranids, or (3) labrids and scarids with vigorous, rapid spawning movements (which could turbulently diffuse gamete clouds) and those with slow movements. Further, egg density does not decline with increasing egg volume among those fishes. Assuming that egg size can affect fertilization success, then sperm limitation seems unlikely to represent a significant problem for pair-spawning T. bifasciatum, probably because mates place their vents close together during gamete release. The situation regarding sperm limitation in other fishes, and effects of environmentally generated water turbulence on it, are less clear. Interspecific variation in the size and content of these fishes' eggs may relate to provisioning of offspring for different larval life-histories.     

Prey nutritional quality and the effectiveness of chemical defenses against tropical reef fishes

Summary Many coral-reef seaweeds and sessile invertebrates produce both secondary chemicals and mineral or fibrous skeletal materials that can reduce their susceptibility to consumers. Although skeletal materials often have been assumed to function as physical defenses, their deterrent effectiveness may derive from their reduction of prey nutritional quality as well as from noxiousness of the skeletal material itself. To test the relative importance of prey nutritional quality and chemical defenses in susceptibility to predation, we offered reef fishes on Guam a choice of artificial foods varying in nutritional quality (4% versus 22% protein) and in secondary chemistry (spanning approximately natural concentration ranges). Field feeding assays were performed with pachydictyol A from the pantropical brown seaweed genus Dictyota, manoalide from the Micronesian sponge Luffariella variabilis, and a brominated diphenyl ether from the Micronesian sponge Dysidea sp. The results indicated that chemical defenses were less effective in high- than in low-quality foods. In paired assays with metabolite-free controls, all three compounds at natural concentrations significantly reduced feeding by reef fishes only in assays using low-quality food, and not in assays with high-quality food. When fishes were offered an array of artificial foods varying in both food quality and metabolite concentration, food quality significantly affected fish feeding in all three cases, while secondary chemistry was significant in only one. Thus differences in nutritional quality, within the natural range among reef organisms, can be comparable to or greater in importance than secondary chemistry in affecting feeding preferences of their consumers. Reduced nutritional quality may be an important selective advantage of producing indigestible structural materials, in addition to their roles as physical support and defense, in coral reef organisms.     

Estimating the role of three mesopredatory fishes in coral reef food webs at Ningaloo Reef,Western Australia

Within the complex food webs that occur on coral reefs, mesopredatory fish consume small-bodied prey and transfer accumulated biomass to other trophic levels. We estimated biomass, growth and mortality rates of three common mesopredators from Ningaloo Reef in Western Australia to calculate their annual turnover rates and potential contribution to local trophic dynamics. Biomass estimates of the serranid Epinephelus rivulatus (4.46 ± 0.76 g m⁻²) were an order of magnitude greater than two smaller-bodied mesopredatory fishes, Pseudochromis fuscus (0.10 ± 0.03 g m⁻²) and Parapercis clathrata (0.23 ± 0.31 g m⁻²). Growth parameters generated from a von Bertalanffy growth function fitted to size-at-age data, however, indicated that mortality rates for the three mesopredators were similar and that 32–55 % of fish survived each year. Consequently, interspecific differences in annual turnover rates among E. rivulatus (1.9 g m⁻² yr⁻¹), Pa. clathrata (0.10 g m⁻² yr⁻¹) and Ps. fuscus (0.07 g m⁻² yr⁻¹) were an artefact of differences in local biomass estimates. The rapid turnover estimates for E. rivulatus suggest this species is an important conduit of energy within the isolated patch reef habitat where it is typically found, while Ps. fuscus and Pa. clathrata channel smaller amounts of energy from specific habitats in the Ningaloo lagoon. Apparent differences in habitat, diet and turnover rates of the three species examined provide an insight into the different roles these species play in coral reef food webs and suggest that life-history traits allow for variability in the local and spatial contribution of these species at Ningaloo Reef. Moreover, calculating turnover rates of a broader suite of fish species from a range of trophic groups will help better define the role of fishes in coral reef trophic dynamics.

     

Potential effects of fishing on cleaning interactions in a tropical reef

We investigated the effect of fishing on the behavior and abundance of the cleaner fish Labroides dimidiatus on the Great Barrier Reef, Australia, by comparing cleaner fish density (underwater visual census) and behavior (focal animal observations) at a fished and a no-take site. Cleaner fish density, as well as the number of individuals and species cleaned, did not differ significantly between sites; however, L. dimidiatus spent more time interacting (physical contact involving feeding bites) with individual clients at the no-take site, indicating that fishing can indirectly affect mutualistic interactions on tropical reefs.     

Behavioral thermoregulatory abilities of tropical coral reef fishes: a comparison with temperate freshwater and marine fishes

Synopsis Eight species in six different families of tropical marine reef fishes from the Indo-West Pacific region (Naso lituratus, Zebrasoma.flavescens, Balistes fuscus, B. vidua, Forcipiger longirostris, Echidna zebra, Cromileptes altivelis, Canthigaster jactator) were tested for ability to thermoregulate behaviorally in electronic shuttleboxes. All of these species preferred mean temperatures between 20 and 30°C, but differed considerably in thermoregulatory precision. All species avoided lethal high or low temperatures (i.e., they did not die during the tests), and some species thermoregulated as precisely as temperate species. Some temperate species prefer higher temperatures (above 30°C) than do these tropical reef species.     

Comparative phylogeography of Atlantic reef fishes indicates both origin and accumulation of diversity in the Caribbean

Background  

Two processes may contribute to the formation of global centers of biodiversity: elevated local speciation rates (the center of origin hypothesis), and greater accumulation of species formed elsewhere (the center of accumulation hypothesis). The relative importance of these processes has long intrigued marine biogeographers but rarely has been tested.