Ecology of a caribbean coral reef. The Porites reef-flat biotope: Part I. Meteorology and hydrography

Observations on certain conditions of the physical environment and plankton ecology of a Caribbean coral reef form the subjects of a two-part study. Here, physical factors are considered, with special attention directed to their influence on the Porites reef-flat biotope. Meteorologic (temperature, precipitation, wind) and hydrographic (temperature, salinity, tide, sea level, current) conditions are examined in order to determine their influence on water movement over the reef and correlation with seasonal variations in plankton abundance. Shoal-water circulation is characterized with reference to patterns of movement, origin, and volume flow. A relationship between wind velocity and direction to volume flow is examined in order to describe the interaction of these parameters. The effects of low tidal exposures and storms on the dominant coral species Porites furcata Lamarck are also examined. Observed mortalities and physical alterations due to these factors are shown to be significant, resulting in relatively rapid modifications of the reef-flat habitat. A chief overall objective of this study is to obtain a quantitative assessment of drifting net plankton crossing the reef-flat environment, and to evaluate its contribution as a food source to the shoal-reef biota. Integration of the physical observations with the plankton ecology will form the subject of a forthcoming publication.     

Molecular genetic evidence for probable reticulate speciation in the coral genus Madracis from a Caribbean fringing reef slope

For many corals, the existence of morphologically distinct yet sympatric populations/species implies reproductive isolation. Conversely, the presence of many intermediate and overlapping morphologies combined with synchronous, mass spawning suggests incomplete reproductive isolation. In Madracis (Scleractinia: Astrocoeniina: Pocilloporidae), high levels of morphological plasticity among the five most commonly recognized species (M. mirabilis, M. senaria, M. decactis, M. formosa and M. pharensis) on Caribbean reefs led us to question species boundaries. Phylogenetic relationships were investigated at the intra-individual, inter-individual and inter-specific levels using the ITS1-5.8S-ITS2 region (ca. 613 bp) of the ribosomal DNA cistron. Inter-specific divergence was ca. 6%, while intra-individual and intra-specific divergences ranged from 0% to 4.9% and 3.3% to 3.5%, respectively. M. senaria and M. mirabilis formed monophyletic groups. M. formosa, M. decactis and M. pharensis formed a paraphyletic complex. High levels of intra-individual and intra-specific ITS polymorphism in the decactis-formosa-pharensis cluster may be the result of very recent speciation within the clade (i.e. maintenance of ancestral polymorphism and incomplete lineage sorting), or the result of repeated introgressive hybridization among the three taxa. Polymorphism parsimony of 89 sites, including nine that showed additivity, revealed a phylogenetic topology more consistent with inter-taxal hybridization. Results are discussed in terms of weak reproductive barriers, and phylogenetic fission and fusion under Veron's model of reticulate speciation in corals. Ecological studies involving Madracis should consider M. decactis, M. formosa and M. pharensis as a complex.     

Effects of ultraviolet radiation and water motion on the reef coral Porites compressa Dana: a flume experiment

The effects of water flow and ultraviolet radiation (UVR, 280–400 nm) on the reef coral Porites compressa Dana were explored in a manipulative flume experiment. The aim of this study was to determine whether this coral responds to changes in the UVR environment by adjusting the tissue concentration of UV-absorbing compounds (mycosporine-like amino acids, MAAs), and to see whether such an acclimation is affected by water flow. Also, calcification rate and chlorophyll-a concentration were measured during the experiment to estimate the potential costs (in the form of slowed growth and/or reduced photosynthetic capacity) to the coral–alga symbiosis of being exposed to UVR and producing MAAs. Branches of P. compressa from a single male colony were exposed to high or low flow (15 cm s⁻¹ and 3 cm s⁻¹, respectively) and ambient or no UVR in an outdoor, continuous-flow seawater system. Chlorophyll-a and MAA concentrations were determined after zero, 3 and 6 weeks of exposure to the experimental conditions. Increase in buoyant weight during the two 3-week periods was used to calculate calcification rate. The presence of UVR had a significant positive effect on total MAA concentration in the P. compressa colonies; however, there were significant interactions present. In colonies exposed to UVR, MAA concentration increased and then decreased to initial levels in high water flow, and increased steadily in low water flow. In colonies receiving no UVR, MAA concentration decreased steadily, declining 23% in 6 weeks. The absence of UVR did not result in higher chlorophyll-a concentrations, but the calcification rate was slightly affected by UVR. This study supports the putative photoprotective role of MAAs in P. compressa, and suggests that the costs of mitigating the effects of ambient UVR are detectable, but they are very small. Received: 29 February 2000 / Accepted: 20 September 2000     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Cleaner-client interactions on a Caribbean reef: influence of correlates of parasitism

We investigated the influence of known correlates of parasitism, namely fish density, body size and social behaviour, on three highly variable aspects of the interactions between cleaning gobies (Elacatinus spp.) and their clients, on a Barbadian coral reef. We specifically considered (1) variability in client visit rate to cleaning stations, (2) cleaning goby preference for specific clients and (3) variation in the time spent cleaning by cleaners. Using phylogenetically independent contrasts, we found that client species that were abundant on the reef visited cleaning stations more often than rarer client species. This could be due to the positive relationship between ectoparasite load and client density, or alternatively may simply reflect the frequency of contact between cleaners and clients. Cleaning gobies spent more time cleaning large-bodied clients, which usually have higher ectoparasite loads, although cleaning goby preference for clients was influenced by none of the correlates of client parasitism. Overall, factors assumed to correlate with ectoparasite load had a limited influence on the variability observed in the interactions between cleaning gobies and their clients. Received: 27 October 1999 / Received in revised form: 11 January 2000 / Accepted: 24 January 2000     

Sexual reproduction of the reef coral Montastreacavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia

Sexual reproduction of the reef-building coral Montastreacavernosa was studied in the Santa Marta area, Caribbean coast of Colombia, from October 1990 to October 1991. The area is subjected to a seasonal upwelling–outwelling regime. From microscopical analysis of tissue sections sampled each lunar month around the full moon, it was confirmed that this is a gonochoric, broadcasting species, with a single gametogenic cycle per year, and a 1:1 sex ratio. Oogenesis began a few weeks after spawning, and extended for 11 mo. The onset of spermatogenesis occurred just before the full moon of June, later than at other localities where sea-water temperatures during the first half of the year are not as low. The amount of reproductive tissue strongly increased in both sexes after July, in association with an increase in sea-water temperature and a decrease in photoperiod. The gonad index of fully mature female colonies was about four times lower than at other localities, perhaps due to the stressful seasonal regime. The male gonad index was lower than that of the female, indicating sex-related differences in the rates of biomass allocation to reproduction. As in other Caribbean localities, spawning occurred after the full moons of August and September. However, there was some evidence of a third spawning episode after the full moon of October, possibly associated with a delay in the occurrence of maximum sea-water temperatures in near-equatorial localities compared to higher latitudes. The probability of cross-fertilization in this species with a gonochoric breeding system and a broadcasting mode of reproduction is favored by its balanced sex ratio, its usually high within-reef zone abundance, and by localized and repeated spawning episodes, synchronized by lunar phase. Received: 19 April 1996 / Accepted: 24 September 1996     

The effects of mating system on male mate choice in a coral reef fish

Summary We show how mate limitation appears to be critical in determining whether or not males exercise mate choice among available females. Thalassoma bifasciatum is a Caribbean reef fish with two distinct mating patterns: group-spawning and pair-spawning. In both mating systems, female fecundity is variable and size dependent, and female availability is high. However, sperm competition among group-spawning males apparently limits the number of effective matings in which a male may engage, whereas territorial pair-spawning males have little or no such limitation. Group-spawning males should be discriminating in their choice of mates and our data confirm this: there is strong evidence for assortative mating in group-spawns, with more large males joining in mating groups around large females. In contrast, pair-spawning males show no indication of mate preferences, and spawn with all females who arrive at their territories.     

A non-motorized dye ejector for visualization of flow in situ and its use with coral reef crinoids

We describe a portable, non-motorized device for delivering a tracer dye into seawater under field conditions. Dye is ejected at a constant flow rate over a period of tens of minutes. The ejector works in a wide range of ambient pressures without external energy requirements. The flow rate is adjusted simply by varying the length of the delivery tube. The dye streams permitted observations of the upcurrent and downcurrent flow regimes for a filter-feeding crinoid (Comanthus bennetti) living at a depth of 8 m on a coral reef. The results indicate that the crinoid may enhance the rate of particle capture by changing the scale of turbulence in the water passing through the mesh of the filtration fan.     

Transmission distance of chemical cues from coral habitats: implications for marine larval settlement in context of reef degradation

The present study (Ishigaki Island, Japan) explored the distance of transmission of chemical cues emitted by live versus dead coral reefs (Exp. 1: High performance liquid chromatography (HPLC) analyses with water sampling station at 0, 1, and 2 km away from the reef) and the potential attraction of these chemical cues by larval fish, crustaceans, and cephalopods (Exp. 2: choice flume experiment conducted on 54 Chromis viridis larvae, 52 Palaemonidae sp larvae, and 16 Sepia latimanus larvae). In the experiment 1, HPLC analyses highlighted that the live coral reef (and not the dead coral reef) produced different and distinct molecules, and some of these molecules could be transported to a distance of at least 2 km from the reef with a reduction of concentration by 14–17-fold. In the experiment 2, C. viridis, Palaemonidae sp, and S. latimanus larvae were significantly attracted by chemical cues from a live coral reef (sampling station: 0 km), but not from a dead coral reef. However, only C. viridis larvae detected the chemical cues until 1 km away from the live coral reef. Overall, our study showed that chemical cues emitted by a live coral reef were transported farthest away in the ocean (at least 2 km) compared to those from a dead coral reef and that fish larvae could detect these cues until 1 km. These results support the assumption of a larval settlement ineffective in degraded coral reefs, which will assist conservationists and reef managers concerned with maintaining biodiversity on reefs that are becoming increasingly degraded.     

Auditory and olfactory abilities of pre-settlement larvae and post-settlement juveniles of a coral reef damselfish (Pisces: Pomacentridae)

The propagules of most species of reef fish are advected from the reef, necessitating a return to reef habitats at the end of the pelagic stage. There is increasing evidence of active attraction to the reef but the sensory abilities of reef fish larvae have not been characterized well enough to fully identify cues. The electrophysiological methods of auditory brainstem response (ABR) and electroolfactogram (EOG) were used to investigate auditory and olfactory abilities of pre- and post-settlement stages of a damselfish, Pomacentrus nagasakiensis (Pisces, Pomacentridae). Audiograms of the two ontogenetic stages were similar. Pre-settlement larvae heard as well as their post-settlement counterparts at all but two of the tested frequencies between 100 Hz and 2,000 Hz. At 100 and 600 Hz, pre-settlement larvae had ABR thresholds 8 dB higher than those of post-settlement juveniles. Both stages were able to detect locally recorded reef sounds. Similarly, no difference in olfactory ability was found between the two ontogenetic stages. Both stages showed olfactory responses to conspecifics as well as L-alanine. Therefore, the auditory and olfactory senses have similar capabilities in both ontogenetic stages. Settlement stage larvae of P. nagasakiensis can hear and smell reef cues but it is unclear as to what extent larvae use these sounds or smells, or both, as cues for locating settlement sites. An erratum to this article can be found at     

Plankton community dynamics of the central Great Barrier Reef Lagoon: Analysis of data from Ikeda et al.

Plankton data collected by Ikeda et al. (1980) from the central region of the Great Barrier Reef, and spanning two years (1976 through 1978) of zooplankton records, have been analyzed extensively for spatial and temporal patterns. Estimates of net zooplankton (including chaetognaths, copepods, and larvaceans) and microzooplankton (juvenile copepods, encompassing nauplii and copepodites, and ciliates) were assessed at three stations across the 60 km lagoon. Temperature, salinity, and chlorophyll a were also measured. A cross-lagoonal gradient was identified in the plankton, concurring with results of related surveys of benthic taxa, such as scleractinian corals, soft corals, macro-algae, fish, sponges, crinoids, etc. Two associations of net zooplankton were identified. The first was associated primarily with the inner lagoon; the second with the outer lagoon. The inshore association was characterized by higher abundances of almost all net zooplankton taxa, particularly chaetognaths, copepods, polychaetes, decapods, and meroplanktonic larvae as well as higher concentrations of chlorophyll a. This inshore association wove back and forth across the lagoon through time, dominating the lagoon entirely during periods of high river discharge, reaching the mid-shelf platform reefs in this region, and sometimes being entirely absent during dry periods. Both seasonal and annual peaks in plankton abundance were generally linked with degree of runoff. Summer/autumn peaks of abundance were evident in chaetognaths, copepods, and larvaceans while annual variation was detected in the former two as well as in chlorophyll a concentrations. Depth stratification was noted in juvenile copepods and chlorophyll a concentrations at the center of the lagoon, with higher abundances recorded in deeper waters. The central Great Barrier Reef lagoon was found to be typical of other tropical coastal waters where plankton community dynamics are controlled primarily by physical factors. We suggest that any substantial changes in river discharge in this area will affect plankton production.A.I.M.S. Contribution No. 242     

The Impact of the Hyperacid Ijen Crater Lake. Part II: A Total Diet Study

In Asembagus (East Java, Indonesia), surface water is contaminated with effluent from the hyperacid Ijen Crater Lake. In a previous study, we reported that food crops irrigated with this water had increased concentrations of various elements. Here, we present a total diet study for adults and 6-year-old children to determine if the mean daily intake of a broad range of elements is safe and adequate. For children, the intake of B, Mg, Mn and V is high with Hazard Quotients (HQ) of 1.1 (B), 1.4 (Mn) and 1.4 (V), respectively (no TDI is available for Mg). For Mn, the daily intake is high due to the consumption of locally produced rice. Drinking water is the main source of B, Mg and V. For adults, the intake of Mg, Mn and V is also high but HQ values are ≤0.7. For both children and adults, the intake of Ca, Zn and particularly Fe is below the RNI (Fe intake is 90% below the RNI for women). It is concluded the mean intake of elements is unbalanced and Fe deficiency is probably the most serious health problem. Toxic effects cannot be fully excluded since deficiency of essential elements such as Ca, Fe and Zn can increase the absorption and retention of various elements.     

Photoinhibition of Symbiodinium spp. within the reef corals Montastraea faveolata and Porites astreoides: implications for coral bleaching

It is speculated that differences in coral bleaching susceptibility may be influenced by the genotype of in hospite Symbiodinium and their differential responses to bleaching stressors. Photoinhibition of photosystem II (PSII), damage to the D1 (psbA) PSII reaction centre protein and production of reactive oxygen species by in hospite Symbiodinium are likely precursors of coral bleaching. In order to assess whether photorepair rates of in hospite Symbiodinium underlie the bleaching susceptibility of their hosts, photoinhibition (net and gross), photoprotection and photorepair rates were assessed in a bleaching-‘tolerant’ coral (P. astreoides) and a bleaching-‘sensitive’ coral (M. faveolata) using non-invasive fluorometric techniques and by blocking de novo synthesis of psbA. Previous studies using such techniques have demonstrated that in vitro Symbiodinium types ‘sensitive’ to bleaching stressors had reduced rates of photorepair relative to ‘tolerant’ Symbiodinum types. Our measurements demonstrated that Symbiodinium in the more bleaching tolerant P. astreoides had higher photorepair rates than Symbiodinium in M. faveolata. Higher repair rates in P. astreoides resulted in lower net photoinhibition relative to M. faveolata, where both corals exhibited similar susceptibility to photodamage (gross photoinhibition). Photoprotective mechanisms were observed in both corals; M. faveolata exhibited higher antennae-bed quenching than P. astreoides at low-light intensities, but at and above light-saturating intensities, which are different for each coral species, P. astreoides displayed more efficient non-photochemical quenching (Stern–Volmer quenching) of chlorophyll fluorescence than M. faveolata. Increased NPQ by P. astreoides at E/E _k ≥ 1 was not driven by antennae-bed quenching. The ability of in hospite Symbiodinium in P. astreoides to mitigate the effects of photoinhibition under high light conditions compared with Symbiodinium in M. faveolata, and their high repair capacity following photoinhibition, may be a key factor to consider in future bleaching studies and may underlie the relative bleaching tolerance of P. astreoides compared to M. faveolata.     

Plankton of the fladen ground during FLEX 76 II. Population dynamics and production of Thysanoessa inermis (Crustacea: Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m^-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m^-2 d^-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m^-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.JOSDAP Contribution No. 50     

Movement patterns of young Caribbean reef sharks, Carcharhinus perezi, at Fernando de Noronha Archipelago, Brazil: the potential of marine protected areas for conservation of a nursery ground

The movement patterns and long-term site-fidelity of primarily juvenile Caribbean reef sharks, Carcharhinus perezi, were investigated using tag-recapture and automated telemetry at an insular nursery area, the Fernando de Noronha Archipelago, Brazil. Of the 143 externally tagged juvenile sharks (<110 cm), 22 (15.3%) were recaptured between 0 and 5 km from the site of tagging after 5–800 days at liberty, suggesting some site-fidelity in young individuals of this species. Site-fidelity and movement patterns of ten juvenile sharks ranging from 78 to 110 cm total length (TL) and one opportunistically captured adult female (224 cm TL) were also investigated for periods of up to 2 years with an array of automated telemetry receivers. Tagging and telemetry data from both inside and outside a marine protected area (MPA) show that shark abundance and activity is greatest along the part of the archipelago’s coastline least disturbed by human activity. Telemetry tracking also showed that juvenile reef sharks demonstrated a high degree of site-fidelity and occupied specific locations along the coast throughout the year, with some evidence of an increase in activity space with ontogeny. Sharks appeared to range more widely at night and there were no seasonal variations in habitat use. Our results suggest that MPAs may be a useful conservation tool to protect young C. perezi and potentially other reef-dwelling carcharhinid sharks during their early life history.     

Concentration fluctuations in a downtown urban area. Part II: analysis of Joint Urban 2003 wind-tunnel measurements

An analysis of concentration time series measured in a boundary-layer wind tunnel at the University of Hamburg is presented. The measurements were conducted with a detailed aerodynamic model of the Oklahoma City (OKC) central business district (CBD) at the scale of 1:300 and were part of the Joint Urban 2003 (JU2003) project. Concentration statistics, as well as concentration probability density (PDF) and exceedance probability (EDF) functions were computed for street- and roof-level sites for three different wind directions. Taking into account the different length scales and wind speeds in the wind-tunnel (WT) and full-scale experiments, dimensionless concentrations and a dimensionless time scale are computed for the comparison with data from the JU2003 full-scale tracer experiments, conducted in OKC in 2003. Using such dimensionless time, the WT time series cover a ~20 times longer time span than the JU2003 full-scale time series, which are analysed in detail in an accompanying, first part of this paper. The WT time series are thus divided into 20 consecutive blocks of equal length and the statistical significance of parameters based on relatively short records is assessed by studying the variability of the concentration statistics and probability functions for the different blocks. In particular at sites closer to the plume edge, the results for the individual blocks vary significantly and at such sites statistics from short records are not very representative. While the location of three sampling sites in the WT closely matched the sites during the full-scale experiments, the prevailing wind directions during the JU2003 releases were not exactly matched. The comparison between full-scale and WT concentration parameters should thus primarily be interpreted in a qualitative rather than direct quantitative sense. Given the differences in mean wind directions and concerns about the representativeness of full-scale concentration statistics, the WT and full-scale results compared well. The 98 percentile concentrations for almost all full-scale releases analyzed are within the scatter of the percentiles observed in the block analysis of the WT time series. Furthermore, the concentration percentiles appear linearly correlated with the fluctuation intensities and the linear relationships determined in the wind tunnel agree well with full-scale results.     

Studies on reef corals. II. Fine structure of planktonic planula larva of Pocillopora damicornis,with emphasis on the aboral epidermis

Tissue and cellular organization of the planula larva of the reef-coral Pocillopora damicornis are detailed ultrastructurally. Emphasis is placed on fine structure of the aboral (=presumptive calicoblast) epidermis. The pre-settling exploratory behavior of the coral larva is discussed. The aboral epidermis consists of a tall columnar epithelium, made up of a heterogeneous cell population-nematocytes, secretory and mucus cells, and flagellated cells. Basally, the columnar epithelium is modified into complexly interdigitating cellular processes, appearing as a discrete histological layer. A-type granules (Jha and Mackie, 1967) indicate neurosecretory activity. The flagellated cell contains a peculiarly comlex flagellar apparatus, including a new intracellular structure, an electron-opaque fibrous cup. It extends across the entire width of the flagellated cell apically, enveloping the flagellar basal bodies and paired transverse accessory rootlets. The flagellum main basal rootlet passes freely through an opening in the base of the rootlet cup. The flagellated cell is here viewed as a sensory cell, as suggested by the corona of microvilli encircling the flagellum, the bundles of microvilli, and extensive areas of tight junctions with adjacent cells. There is no evidence in the gastrodermis or epidermis of the planktonic larva for pre-settling accumulation of calcareous skeletal material. Vesicles containing a highly oriented fibrous material may represent sites of skeletal organic-matrix precusor buildup.Contribution no. 451. Hawaii Institute of Marine Biology, Kaneohe, Hawaii.     

The phenology of a Neotropical ant assemblage: evidence for continuous and overlapping reproduction

Reproductive phenologies reflect the interaction between the mating system of a taxon and the local environment. Ant colonies reproduce and disperse via the flights of winged alates. Few data exist on the reproductive phenologies of ant assemblages. Here we analyze the reproductive phenologies of 81 common ant species from 23,182 individuals collected over 3 years on Barro Colorado Island, Panama (BCI). Species ranged from highly synchronous to continuous fliers, but showed a median flight duration of at least 8 of 13 lunar months. In two statistical analyses (variance ratio test and Spearman rank correlations), 84% (16 of 19) of ant genera had species trending toward positively associated phenologies, more than expected by chance (P<0.00036 by a binomial test). Thus, there was little evidence for the hypothesis that competition for limiting resources staggers congeneric flights and ultimately promotes reproductive isolation. On the contrary, the timing of reproduction, and its synchrony, tended to be conserved within genera and subfamilies. These results closely match phenological studies of plant assemblages. The continuous reproduction and small colony size of many species in this study suggest that the female calling syndrome, a poorly documented mating system in ants, may be common on BCI.