Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Indirect effects of a key ecosystem engineer alter survival and growth of foundation coral species

Stegastes nigricans, a "farmerfish" that cultivates algal turf and defends territories from grazers and other intruders, can affect coral indirectly due to increased competition with farmed algal turf and/or reduced predation resulting from territorial aggression directed at corallivores. To investigate the indirect effects of this key ecosystem engineer on coral mortality and growth, we transplanted caged and exposed fragments of four coral species to patch reefs in French Polynesia on which we manipulated the presence of S. nigricans and turf, and to reefs naturally devoid of S. nigricans. Reef access was two to four times higher for herbivorous fishes, and two times higher for corallivorous fishes, when S. nigricans was removed, indicating that reef access is reduced for two important guilds of fishes when S. nigricans is present. Stegastes' territoriality indirectly benefited delicate acroporids (Montipora floweri and Acropora striata), yielding a twofold to fivefold reduction in skeletal loss due to lower predation frequencies in the presence of S. nigricans. Three corals, A. striata, M. floweri, and especially Porites australiensis, suffered mortality due to overgrowth significantly more frequently in the presence of farmed turf, but Pocillopora verrucosa did not. Algal abundance predicted the frequency of overgrowth for only A. striata and P. australiensis. M. floweri were more likely to be overgrown when exposed (uncaged) in the presence of S. nigricans, suggesting an interaction modification, in this case that initial predation increased susceptibility to competition with turf. In this community, the presence of S. nigricans may increase algal overgrowth of massive Porites by facilitating its turf competitors and simultaneously reduce predation of branching corals through territorial exclusion of corallivores. These indirect interactions may underlie previously documented community transitions from disturbance-resistant massive coral to recovering branching corals within S. nigricans territories.     

Bioerosion of coral reefs by two Hawaiian parrotfishes: species, size differences and fishery implications

Parrotfishes can be significant bioeroders and sediment producers on coral reefs. We quantified the bioerosion rates of two similarly sized Hawaiian parrotfishes with two different feeding modes (Scarus rubroviolaceus—a scraper and Chlorurus perspicillatus—an excavator). The results showed that feeding modes did not affect bioerosion rates but that bioerosion rates were size dependent, with largest individuals (S. rubroviolaceus 45–54 cm FL) bioeroding up to 380 ± 67 kg ind⁻¹ year⁻¹. The size for onset of bioerosion capabilities for both species was 15 cm. Grazing by the two species consumed 60% of the carbonate production of the fore reef area, suggesting that large parrotfishes in Hawaii are ecologically important bioeroders. As individual large S. rubroviolaceus contributed disproportionately more to bioerosion and sediment production than the equivalent biomass of smaller conspecifics, management strategies designed to retain normal reef bioerosion rates should seek to preserve the historical size structure of S. rubroviolaceus populations and to especially protect the larger size classes.     

Illumination and the coral community beneath tabular Acropora species

Light profiles beneath the tabular coral species Acropora cytherea and A. hyacinthus were examined at Rib and Broadhurst Reefs, central Great Barrier Reef, in November 1980. They show a strongly decreasing illumination gradient towards the central stem. Beneath tables at 10m deep, which receive about 7.0 mW cm^–2 at solar noon, substrate irradiance falls to a minimum of about 5% ambient or to about 0.4 mW cm^–2. Thus, from previously reported compensation values, most sub-table substrate may receive above-compensation irradiance. Illumination beneath tables screened by foil is significantly lower. The coral communities beneath tables of these two species plus those of A. subulata and A. clathrata support an average 26 colonies m^–2, at a cover of 40%, nearly identical to adjacent, unshaded quadrats. Species richness is likewise very similar. Species richness and colony density beneath the tables increase inwards from the perimeter of the shaded areas, decreasing only near the central stems. Dark adaptation and reduced competitive and grazing pressure are suggested explanations for the latter. It is concluded that shading does not provide a significant competitive advantage for Acropora spp. tables at 10 m deep.     

Structural investigations on the mucus from six species of coral

The chemical composition of the mucus from three hard corals (Acropora formosa, Pachyseris speciosa and Fungia fungites) and three soft corals (Sarcophyton sp., Lemnalia sp., and Cespitularia sp.) collected on the Great Barrier Reef (1982–1985) was determined. Significant variation exists in the composition and structure of the six mucus samples, indicating the absence of a common structure for coral mucus. In all cases protein and/or carbohydrate polymers are the major components of the mucus, and lipids are present only in small amounts. The glycose composition varied between species, with fucose (F. fungites and Lemnalia sp.), arabinose (A. formosa), galactose (P. speciosa) and N-acetyl glucosamine (Sarcophyton sp.) being present in high concentrations. With the exception of Sarcophyton sp. and Lemnalia sp., all mucus samples were acidic and contained significant sulphate but no uronic or sialic acids. The amino acid composition of the mucus samples was not unusual, apart from A. formosa, which contained a high percentage of serine and threonine, and F. fungites, which had high levels of glutamic acid present.     

Hybridization within the species complex of the scleractinan coral Montastraea annularis

The morphologically variable reef coral previously known as Montastraea annularis (Ellis and Solander, 1786) has recently been separated into three species based on differences in morphology, behavior, allele frequencies and some life-history traits of Panamanian specimens. To further investigate the proposed reclassification and its conformity to the biological species concept we conducted reciprocal intra- and inter- specific fertilization experiments with gametes from each of the three species on Florida reefs. With one exception, self-fertilization rates were very low or zero. Within-species crosses resulted in production of planulae, as did all inter-species (hybrid) crosses, but there was much variation in fertilization success within each type of cross. In an experiment with separated gametes, hybrid crosses between M. annularis and M. franksi produced more larvae than within-species crosses for each species. Hybridization crosses between M. faveolata and the other two species produced fewer larvae than did within- M. faveolata crosses in the experiment with separated gametes, but many larvae resulted when the hybridizations were performed by mixing entire gamete bundles. Additional observations showed that M. franksi had 20% larger eggs and fewer eggs per gamete bundle than did the other two species and that it consistently spawned 1 to 1.5 h before the others, a potential temporal barrier to hybridization. These results indicate that there is no inherent pre-zygotic barrier to cross-fertilization among the three morphological species, although post-zygotic survival and fertility remain to be determined. The adherence of the proposed reclassification to the biological species concept requires further examination. Received: 16 April 1997 / Accepted: 17 June 1997     

Sexual reproduction of three coral species from the Mexican South Pacific

Our understanding of the reproductive biology of corals from the Mexican southeastern Pacific is limited, and consequently, the role of reproduction in structuring coral communities is unclear. As a first attempt to understand the importance of sexual reproduction in structuring and maintaining of the coral communities from this region, we documented the reproductive cycles over 2 years (2003–2004) in three main reef-building corals species in the region. Pocillopora damicornis was shown to be hermaphroditic with asynchronous gamete development observed only in 2004; P. gigantea was characterized as both gonochoric and cosexually hermaphroditic. Absence of mature gametes was documented in both species, and an observation may be attributed to the 2003 El Niño Southern Oscillation event, which may have inhibited reproductive maturation via thermal stress. Porites panamensis was gonochoric with asynchronous development, and planulae were generally brooded. The presence of mature gametes and planulae in P. panamensis polyps suggests that this species is an important contributor to local and, likely, regional recruitment of this species. Further research should seek to identify important source populations for these coral recruits and document the exchange of larvae between coral populations of the Mexican Pacific.     

Availability of two forms of dissolved nitrogen to the coral Pocillopora damicornis and its symbiotic zooxanthellae

The relative contribution of dissolved nitrogen (ammonium and dissolved free amino acids DFAAs) to the nitrogen budget of the reef-building coral Pocillopora damicornis was assessed for colonies growing on control and ammonium-enriched reefs at One Tree Island (southern Great Barrier Reef) during the ENCORE (Enrichment of Nutrient on Coral Reef; 1993 to 1996) project. P. damicornis acquired ammonium at rates of between 5.1 and 91.8 nmol N cm⁻² h⁻¹ which were not affected by nutrient treatment except in the case of one morph. In this case, uptake rates decreased from 80.5 to 42.8 nmol cm⁻² h⁻¹ (P < 0.05) on exposure to elevated ammonium over 12 mo. The presence or absence of light during measurement did not influence the uptake of ammonium ions. Nitrogen budgets revealed that the uptake of ammonium from concentrations of 0.11 to 0.13 μM could completely satisfy the demand of growing P. damicornis for new nitrogen. P. damicornis also took up DFAAs at rates ranging from 4.9 to 9.8 nmol N cm⁻² h⁻¹. These rates were higher in the dark than in the light (9.0 vs 5.1 nmol m⁻² h⁻¹, P < 0.001). Uptake rates were highest for the amino acids serine, arginine and alanine, and lowest for tyrosine. DFAA concentrations within the ENCORE microatolls that received ammonium were undetectable, whereas they ranged up to 100 nM within the control microatolls. The contribution of DFAAs to the nitrogen budget of P. damicornis constituted only a small fraction of the nitrogen potentially contributed by ammonium under field conditions. Even at the highest field concentrations measured during this study, DFAAs could contribute only ≃11.3% of the nitrogen demand of P.␣damicornis. This contribution, however, may be an important source of nitrogen when other sources such as ammonium are scarce or during periods when high concentrations of DFAAs become sporadically available (e.g. cell breakage during fish-grazing). Received: 22 April 1998 / Accepted: 3 November 1998     

Comparing ecosystem engineering efficiency of two plant species with contrasting growth strategies

Many ecosystems are greatly affected by ecosystem engineering, such as coastal salt marshes, where macrophytes trap sediment by reducing hydrodynamic energy. Nevertheless, little is known about the costs and benefits that are imposed on engineering species by the traits that underlie their ecosystem engineering capacity. We addressed this topic by comparing ecosystem engineering efficiency defined as the benefit-cost ratio per unit of biomass investment for two species from the intertidal habitat: the stiff grass Spartina anglica and the flexible grass Puccinellia maritima. These species were selected for their ability to modify their habitat by trapping large quantities of sediment despite their contrasting growth form. On a biomass basis, dissipation of hydrodynamic energy from waves (a proxy for benefits associated with ecosystem engineering capability as it relates to the sediment trapping capability) was strikingly similar for both salt marsh species, indicating that both species are equally effective in modifying their habitat. The drag forces per unit biomass (a proxy for costs associated with ecosystem engineering ability as it relates to the requirements on tissue construction and shoot anchoring to prevent breaking and/or washing away) were slightly higher in the species with flexible shoots. As a result, stiff Spartina vegetation had slightly higher ecosystem engineering efficiency, due to lower engineering costs rather than to a higher engineering effect. Thus, Spartina is a slightly more efficient rather than a more effective ecosystem engineer. Ecosystem engineering efficiency was found to be a species-specific characteristic, independent of vegetation density and relatively constant in space. Analyzing ecosystem engineering by quantifying trade-offs offers a useful way toward developing a better understanding of different engineering strategies.     

Temporal and spatial variability in coral recruitment on two Indonesian coral reefs: consistently lower recruitment to a degraded reef

Corals are the primary reef-building organisms, therefore it is key to understand their recruitment patterns for effective reef management. Coral recruitment rates and juvenile coral abundance were recorded in the Wakatobi National Marine Park, Indonesia, on two reefs (Sampela and Hoga) with different levels of environmental degradation (12.5 vs. 44 % coral cover with high and low sedimentation rates, respectively) to examine consistencies in recruitment patterns between years and seasons. Recruitment was measured on multiple panels at two sites on each reef (6–7 m depth) and cleared areas of natural reef. Although coral recruitment was twofold higher in 2008–2009 than in 2007–2008, and seasonal differences were identified, consistent significant differences in recruitment rates were found between the two reefs even though they are separated by only ~1.5 km. Recruitment rates and juvenile abundance were lower on the more degraded reef. These patterns are likely a consequence of differential pre- and post-settlement mortality as a result of the high sedimentation rates and degraded conditions and possibly reduced larval supply.     

Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef

Following observations of mass spawning of hermatypic corals on the Great Barrier Reef in 1981 and 1982, spawning dates were successfully predicted and documented at five reefs on the Central and Northern Great Barrier Reef in 1983. During the predicted times, 105 species from 36 genera and 11 families were observed to spawn. Of these, 15 species were shown to have an annual gametogenic cycle. All but two of the species observed during mass spawnings shed gametes which underwent external fertilization and development. Synchronous spawning was observed both within and between the five reefs studied, which were separated by as much as 5° of latitude (500 km) or almost a quarter of the length of the Great Barrier Reef. The mass spawning of corals took place on only a few nights of the year, between the full and lastquarter moon in late spring. Maturation of gametes coincided with rapidly rising spring sea temperatures. Lunar and diel cycles may provide cues for the synchronization of gamete release in these species. The hour and night on which the greatest number of species and individuals spawned coincided with low-amplitude tides. Multispecific synchronous spawning, or mass spawning, of scleractinian and some alcyonacean corals represents a phenomenon which is, so far, unique in both marine and terrestrial communities.     

Hidden genetic diversity in a key model species of coral

A total of 43 colonies of the scleractinian coral Pocillopora damicornis from lagoonal and reef slope sites in the western Indian Ocean (WIO) region were genetically characterised at one nuclear and two mitochondrial sequence markers and six microsatellite loci. Both mitochondrial and microsatellite data support the existence of two reciprocally monophyletic clusters (F- and NF-types) and provide evidence of the existence of two cryptic species of P. damicornis on reefs in WIO region and put current morphological delineation and geographical boundaries of P. damicornis and Pocillopora molokensis into question. The results add to ongoing studies on the phylogeny and phylogeography within the genus Pocillopora, which all point towards a range of unresolved morphological and molecular species boundaries. Nuclear phylogenies derived from the present and previously published sequences show evidence for incomplete lineage sorting and/or introgressive hybridisation between Pocillopora morphospecies. However, the two WIO types largely remain in separate clusters, further supporting the theory that these represent two different species.     

Large ontogenetic declines in intra-crown leaf area index in two temperate deciduous tree species

The widespread occurrence of age-related changes in leaf morphology and allocation suggests that the leaf area index of individual trees (intra-crown LAI) may decline late in ontogeny. We used direct, within-canopy measurements to quantify the LAI of canopy trees with exposed crowns of two temperate deciduous species. Intra-crown LAI declined from approximately 7 to 4 in Acer saccharum, and from approximately 9.5 to 6.5 in Betula alleghaniensis, as tree size increased (from 15 to 72 cm diameter at breast height [dbh]). For A. saccharum, age (which varied from 30 to 160 years) was a significantly better predictor of LAI decline than dbh. We also modeled the effect of ontogenetic declines in LAI on understory light availability and found that light transmission increases significantly as canopy trees grow and mature. Our results thus suggest that gradual declines in LAI with tree age may play an important and overlooked role in contributing to the heterogeneity of sub-canopy light regimes in mature forests.     

Testing hypotheses of exponential growth and size-dependent molting rate in two copepod species

Individual copepodids from nature of the lipidstoring, monocyclic, diapausing Calanus glacialis, and from nature and laboratory culture of the non-storing, polycyclic Eurytemora herdmani (both collected new Halifax, Nova Scotia in 1987 and 1988) were reared in excess food at ca. 3 and 10 °C. Soon after molts, prosome lengths and weights [total dry wt (TW) of E. herdmani; structural wt (SW) and estimated oil-sac wt (OSW) of C. glacialis] were measured. Stage durations were close to published temperature-dependent predictions; C. glacialis (almost all females) did not enter resting stages. Growth of body length was linear and of body weight (TW or SW) was exponential, with no sexual difference in E. herdmani. There were mixed effects of sizes on stage durations of individuals: weakly positive at 3 °C (but not significant at ca. 10°C) for length and SW of C. glacialis and generally weakly negative for E. herdmani, except for TW at 10 °C. Body condition (residuals of log SW vs log length) of C. glacialis at ca. 10°C, unlike length or SW, was negatively related to stage duration. There was no relationship between length-corrected SW and OSW in C. glacialis. The various results appear to suggest that health was more important than allometric constraints on growth rates of individuals. However, copepods reared at temperatures very different from those previously experienced may show long-term adjustments of size, whereas development rates respond immediately. If so, only the weakly positive effects on stage durations of length of C. glacialis at ca. 3 °C and of TW of E. herdmani at ca. 10 °C may illustrate expectations under stable temperatures and adequate food in nature.     

Estimating the footprint of pollution on coral reefs with models of species turnover

Ecological communities typically change along gradients of human impact, although it is difficult to estimate the footprint of impacts for diffuse threats such as pollution. We developed a joint model (i.e., one that includes multiple species and their interactions with each other and environmental covariates) of benthic habitats on lagoonal coral reefs and used it to infer change in benthic composition along a gradient of distance from logging operations. The model estimated both changes in abundances of benthic groups and their compositional turnover, a type of beta diversity. We used the model to predict the footprint of turbidity impacts from past and recent logging. Benthic communities far from logging were dominated by branching corals, whereas communities close to logging had higher cover of dead coral, massive corals, and soft sediment. Recent impacts were predicted to be small relative to the extensive impacts of past logging because recent logging has occurred far from lagoonal reefs. Our model can be used more generally to estimate the footprint of human impacts on ecosystems and evaluate the benefits of conservation actions for ecosystems.     

Coral weight increment in situ. A new method to determine coral growth

An underwater weighing apparatus is described. This consits basically of a float, the buoyancy of which is neutralized by a known amount of lead weights. The standard error of the method is ±0.1g. The entire weighing procedure takes place on the reef using SCUBA diving gear. The advantages are that the animals are not harmed by experimental treatment, and that the pure calcium-carbonate increment is registrated. Some results obtained by this method show growth of Montastrea annularis and Madracis mirabilis.     

Variation in benthic communities of eastern Caribbean coral reefs in relation to surface sediment composition

The effects of sedimentation on coral reefs are commonly studied at local scales, but larger-scale patterns have been elusive, making it difficult to determine the role of sedimentation in region-wide changes in these ecosystems. We examined the relationships between characteristics of reef-associated surface sediment and benthic composition of 22 reefs around 11 islands of the eastern Caribbean. The terrigenous fraction in surface sediment increased with proximity to a clear source of sediment input. The percent cover of live coral, macroalgae, and turf algae decreased with higher terrigenous sediment fraction, while sponge cover increased. Sites with sediment containing high and low terrigenous fraction differed in coral species assemblages. In particular, the cover of Montastraea annularis complex decreased with increasing terrigenous sediment fraction. The proportion of fine-grained sediment had no effect on benthic composition. These results suggest that sedimentation may play a role in shaping coral reef communities at a regional scale.     

A comparative study of feeding and growth in two coexisting species of carnivorous gastropods

The effects of variable food supply on growth rate and feeding behaviour in two coexisting intertidal gastropods were examined through field and laboratory experiments involving mark and recapture procedures, growth experiments under controlled food supply and video observations of feeding. All cohorts of both Japeuthria species grew most in summer, with the highest growth rate being achieved by the youngest cohort of both sexes. In contrast, almost all cohorts (except the youngest one) showed zero to negative growth in winter. The growth rates of Japeuthria species clearly varied depending on feeding frequencies and the initial size of individuals. Females of both species had a significantly higher growth rate than males, though the difference became less marked with decreasing food supply. The total time spent on feeding and the length of the first feeding bout generally increased as the duration of the interval between meals was increased across treatments. From both the field growth experiment and the observations of feeding behaviour it is clear that J. ferrea had a higher frequency of feeding than J. cingulata in the field; 1- to 3-d intervals of feeding frequency in the former and 3- to 6-d intervals in the latter. Most notably, the two species showed a reversal in growth rates with changing conditions of food supply. When food was relatively abundant J. ferrea achieved better growth than J. cingulata, but J. cingulata grew better than J. ferrea under low food supply. In view of the fact that J. ferrea seems to be competitively superior to J. cingulata under normal levels, the present observation that J. cingulata can perform better than J. ferrea when food availability is low gives an important advantage to the latter in terms of its coexistence with the former. Received: 9 March 1999 / Accepted: 8 October 1999