Tale of two colors: <Emphasis Type="Italic">Cladopsammia gracilis</Emphasis> (Dendrophylliidae) color morphs distinguished also by their genetics and ecology

Cladopsammia gracilis (Dendrophylliidae), an ahermatypic coral inhabits the northern Red Sea. Two color morphs (pink and orange) are found aggregated in caves devoid of hermatypic corals, associated with crustose coralline algae (CCA). Sequencing the rDNA ITS region revealed a separate clustering of members of each color morph. Both morphs grow in shallow waters, with orange corals limited to the upper 4 m, while some pink coral aggregates thrive deeper than 30 m. Planulae were released between June and December. Pink planulae treated with antibiotics and exposed at different intervals to CCA, were competent and metamorphosed even 110 days after release. Maximal competency period for orange planulae was 70 days. All planulae were enhanced to metamorphose in presence of CCA. The mean age at metamorphosis of pink and orange planulae treated with CCA differed significantly. Most orange planulae settled directly on the CCA while most pink planulae settled on the wall of the experiment vial. The morphs differed significantly in the calyx cross-section area of primary polyps. Despite being considered a single species according to skeletal based taxonomy, the significant ecological and molecular differences between pink and orange C. gracilis specimens suggest that they may belong to separate species.     

Ecological and physiological differences between two colour morphs of the coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus) is a ubiquitous branching coral found throughout the Indo-Pacific region. Like many other species of coral, P. damicornis displays a large range of morphologies. At One Tree Island, it occurs as two distinct morphs that are easily distinguished by the presence or absence of pink pigmentation. The two colour morphs of P. damicornis were found to differ in their distribution and abundance in the One Tree Island Lagoon. The brown morph was more abudant than the pink morph in the shallows (<1 m),whereas the pink morph was more abundant at deeper sites (>3 m). The two morphs also differed physiologically. The brown morph tended to have a greater calcification rate than the pink morph, regardless of environmental conditions. However, the difference in the calcification rate between the two morphs became non-significant under shaded conditions (5% full sunlight), indicating some degree of physiological plasticity of the morphs. The pink colour in P. damicornis was due to a hydrophilic pigment with a major peak absorbance at 560 nm. The expression of pink pigment had both genetic and phenotypic components. The brown morph has a reduced genetic capacity to express the pigment relative to the pink morph. On the other hand, pigment expression could be induced by light in the pink morph. Although genetic differences ultimately determine the differences between the two morphs of P. damicornis, the extent of pigment expression is under some degree of environmental influence.     

Biochemical-genetic and ecological evidence that red/brown individuals of the anemone Actinia equina comprise two morphs in Britain

A study of the sea anemone Actinia equina L. displaying red to brown column coloration was conducted on British shores in 1981 and 1982 in order to assess the ecological significance of discontinuous variation in pedal disc colour. Two ecologically distinct morphs have been revealed. One is characterised by having a red/pink pedal disc, and by being homozygous slow or heterozygous at a hexokinase locus, whilst the other has a grey/green disc and is homozygous fast. The former is relatively more common on the upper-mid shore and on vertical rock faces, whereas individuals of the latter occur more frequently at lower levels, and are relatively more abundant on horizontal surfaces. Correlated with this is a dichotomy in the strength of attachment of the disc to the substratum; anemones having red/pink discs adhere relatively more firmly. Evidence from brood size analysis suggest that the morphs are subject to differential selection; the commoner one at many site examined had a larger median brood size. At present, none of our data on gene-enzyme variation suggests that red to brown anemones in Britain comprise more than one species. Nevertheless, it is argued that local speciation is potentially likely to occur as a result of disruptive selection.     

Genetic connectivity among color morphs and Pacific archipelagos for the flame angelfish, <Emphasis Type="Italic">Centropyge loriculus</Emphasis>

Color variation is used in taxonomic classification of reef fishes, but it may not reliably indicate evolutionary divergence. In the central Pacific, there are three color morphs of the flame angelfish, Centropyge loriculus: a red morph that occurs primarily in the Hawaiian archipelago, the endemic Marquesan color morph with reduced black markings, and an orange morph that occurs throughout the rest of Oceania. The red and orange morphs co-occur at Johnston Atoll (1,300 km south of Hawai’i), but intermediate forms have not been reported. To determine whether the three color morphs represent distinct evolutionary lineages, we compared 641 base pairs of mitochondrial cytochrome b. Forty-one closely related haplotypes were observed in 116 individuals. Analysis of molecular variance (AMOVA) indicated no significant genetic structure among color morphs (Φ_ST = 0.011, P = 0.147). Likewise, there was no significant pairwise structure between sampling locations, separated by up to 5,700 km, after a Bonferroni correction (Φ_ST = 0.000–0.080, P = 0.0130–0.999). Genetic studies in conjunction with larval distribution data indicate that Centropyge species are highly dispersive. While there is a strong geographic component to the distribution of color morphs in C. loriculus, we find no evidence for corresponding genetic partitioning. We do not rule out an adaptive role for color differentiation, but our data do not support emerging species.     

Variation in the genetic composition of coral (Pocillopora damicornis and Acropora palifera) populations from different reef habitats

The genetic structure of populations of the corals Pocillopora damicornis and Acropora palifera was examined in three habitats at One Tree Island during March and April 1993, using electrophoretically detectable variation at six allozyme loci. There were significant genetic differences among populations of P. damicornis within each of the reef crest, lagoon and microatoll habitats. The level of differentiation among populations was similar in each of the habitats. Differences between populations of P. damicornis from lagoon and microatolls were no greater than that within habitats, but genetic differentiation of these from crest populations was much higher. There was no difference in the genetic composition of A. palifera populations within or between the lagoon and microatolls, the only habitats where this species was found. Both coral species had observed:expected (G _O:G_E) genotypic diversity rations >0.80, indicating predominantly sexual reproduction. These data, the high genotype diversity and general conformance of genotype frequencies to those expected under conditions of Hardy-Weinberg, suggested panmixis at each site. The high degree of sexual reproduction in the P. damicornis populations is unusual for a species where asexual reproduction has been the dominant mode of reproduction reported to date. Gene flow in both species was considerable between the lagoon and the closed microatolls. The genetic differences between populations of P. damicornis in these habitats and the reef crest may reflect the relative isolation of all populations within the closed One Tree Lagoon from those outside. However, local currents appear to offer effective means of dispersal between the habitats, suggesting that the genetic differences result from natural selection in the different environments within One Tree Lagoon and the reef crest.     

Resource-associated divergence in the arctic marine amphipod<Emphasis Type="Italic"> Paramphithoe hystrix</Emphasis>

Host specialization has played an important role in the speciation of groups such as herbivorous insects and parasitic invertebrates. In this study, we provide evidence for its role in the diversification of the amphipod Paramphithoe hystrix Ross 1835 in Arctic marine waters. This species lives and feeds on the tissues of varied invertebrate hosts that are much larger than itself, including poriferans, hydrozoans and echinoderms, acting as a 'micropredator'. We examined the genetic structure of P. hystrix at sites in the Canadian Arctic where it is represented by a white form found on the light-coloured sea star Solaster endeca, a red form on the pink soft coral Gersemia rubriformis and by a spotted form with an unidentified host. These phenotypes occur in microallopatry, sometimes occupying alternate hosts just a few metres apart. Although their variation in host associates might have arisen from ingested host tissue, our results indicate that these morphs are genetically distinct. Mitochondrial (cytochrome c oxidase I) and nuclear (28S ribosomal RNA) gene sequences, as well as allozyme data all show strong genetic divergence between the three colour morphs, demonstrating their reproductive isolation. These results suggest that P. hystrix is a complex of at least three species, which may have arisen as a result of disruptive selection following host switches. Moreover, the depths of genetic divergence indicate that diversification of this complex was complete prior to the Pleistocene.Communicated by R.J. Thompson, St. John's     

Endemism and Regional Color and Genetic Differences in Five Putatively Cosmopolitan Reef Fishes

Abstract:  Endemism is thought to be relatively rare in marine systems due to the lack of allopatric barriers and the potential for long-distance colonization via pelagic larval dispersal. Although many species of coral reef fishes exhibit regionally restricted color variants that are suggestive of regional endemism, such variation is typically ascribed to intraspecific variation. We examined the genetic structure in 5 putatively monospecific fishes from the Indo-West Pacific ( Amphiprion melanopus , Chrysiptera talboti , and Pomacentrus moluccensis [Pomacentridae] and Cirrhilabrus punctatus , and Labroides dimidiatus [Labridae]) that express regional color variation unique to this area. Mitochondrial-control-region sequence analysis showed shallow to deep genetic divergence in all 5 species (sequence divergence 2–17%), with clades concordant with regional color variation. These results were partially supported by nuclear RAG2 data. An analysis of molecular variation (AMOVA) mirrored the phylogenetic results; Φ_ST values ranged from 0.91 to 0.7, indicating high levels of geographic partitioning of genetic variation. Concordance of genetics and phenotype demonstrate the genetic uniqueness of southwestern Pacific color variants, indicating that these populations are at a minimum distinct evolutionarily significant units and perhaps distinct regionally endemic species. Our results indicate that the alpha biodiversity of the southwestern Pacific is likely underestimated even in well-studied groups, such as reef fishes, and that regional endemism may be more common in tropical marine systems than previously thought.     

The hydrozoan coral<Emphasis Type="Italic"> Millepora dichotoma</Emphasis>: speciation or phenotypic plasticity?

This article describes ecological and biological differences between two morphs of the Red Sea fire coral Millepora dichotoma. The species is divided into two main morphs: branching and encrusting, which were found to differ both in color and morphology. Each morph has two or more sub-morphs. A total of 372 M. dichotoma colonies were examined in a census at two study sites in the Gulf of Elat. Colony size and abundance of the two morphs were found to differ significantly between sites. Experimental examination of each morph's morphological plasticity revealed different growth rates and difference in growth plasticity between the branching and the encrusting morph. Most of the fragments from the branching colonies (94%) attached to experimentally placed Plexiglas substrate, compared with much less attachment by the encrusting fragments (11%). The growth form of the branching morph on the Plexiglas switched to encrusting, spreading over and covering the substrate. When the new encrusting colony reached the edge of this substrate, it started to produce tips, and returned to growth in the classic branching form. The encrusting morph did not change its growth form. Following attachment of the original fragments of the branching morph to the substrate, 8.1% of them produced new tips. When the original branches were removed, after converting to encrusting growth form, 19% of the fragments produced new tips. The capsule size of nematocysts of the two morphs was also significantly different (t-test, P<0.05). Molecular data (ITS region) clearly demonstrate that these two M. dichotoma morphs differ considerably. Molecular evidence (srRNA) from the symbiotic zooxanthellae also shows a different pattern of clades in the hosts. The ecological, biological and molecular data thus attest to the two morphs being distinguishable. Contrary to previous reports, we consequently suggest that the two morphs of M. dichotoma found in the Gulf of Elat are actually two distinct species.Communicated by O. Kinne, Oldendorf/Luhe     

Morphological variation and phenotypic plasticity of buoyancy in the macroalga Turbinaria ornata across a barrier reef

Many aspects of morphology of benthic algae (length, surface area-to-volume ratio, and blade undulation) are plastic traits that vary in response to physical factors (such as light or water flow environment). This study examines whether frond buoyancy is a plastic trait, and whether differences in morphology including buoyancy affect the potential persistence of macroalgae in habitats characterized by different water flow regimes. Fronds of the tropical alga Turbinaria ornata in protected backreef environments in Moorea, French Polynesia possess pneumatocysts (gas-filled floats) and experience positive buoyant forces, whereas fronds in wave-exposed forereef sites either lack pneumatocysts entirely or have very small, rudimentary pneumatocysts and experience negative buoyant forces. Forereef fronds transplanted to the backreef developed pneumatocysts and experienced increased buoyant force indicating that buoyancy is a phenotypically plastic trait in T. ornata. In comparing the potential for dislodgement by drag, drag was greater on forereef fronds at low flow speeds as these fronds were stiffer and did not bend over at low flow speeds and therefore were less streamlined in the flow than backreef algae, which bent easily. The environmental stress factor (ESF) (a measure of the likelihood of detachment for a frond in its habitat) was higher for forereef than backreef fronds at all flow speeds. When examined with respect to the flow velocities likely in their respective habitats however, the chance of detachment for backreef and forereef was similar. Neither backreef nor forereef fronds were predicted to break under normal, non-storm conditions, but both were predicted to break in storms. Strong forereef morphologies are well suited to habitats characterized by rapid flow, whereas the weaker, buoyant, tall backreef fronds are well suited to habitats where crowding and shading is common but hydrodynamic forces are low.     

Genetic relationships of the colour morphs of Acanthochromis polyacanthus (Pomacentridae) on the northern Great Barrier Reef

Acanthochromis polyacanthus is an unusual tropical damselfish because it lacks pelagic larvae and has lost the capacity for broad-scale dispersal among coral reefs. On the modern Great Barrier Reef, this fish has multiple colour morphs and we have previously described steep genetic clines in the central region between three colour morphs. A similar transition of colour morphs has been found in the far northern region (Latitude 11°S), where at least five colour morphs and intergrading hybrids were found during a survey of 14 locations on nine adjacent reefs. On two of these reefs, monochromatic white and bicoloured morphs coexist, partitioning the reef slope vertically when sympatric, but occupying the full depth range elsewhere. In areas of overlap, they interbreed, producing hybrids of intermediate colour. Although the discriminatory power of our genetic analysis was weakened by the low level of variability in these populations (only four polymorphic loci), allozyme electrophoresis revealed significant genetic differentiation within and among reefs. There was neither simple linkage between colour and protein variation, nor correlation between genetic similarity and proximity of sampled populations. We suggest that the observed chaotic structure is the result of differences among the founding populations, although the two colour-based stocks have been homogenized through a long history of introgression during multiple episodes of isolation and secondary contact. Despite genetic mixing, phenotypic diversity (i.e. colour morphs) seems to have been preserved in these populations by sexual selection, perhaps reinforced by natural selection. The long-term stability of this outcome, however, is suggested by the presence of unique morphs on two reefs that are detached from the continental shelf and surrounded by very deep water. The strong identity of these populations can be attributed to long occupation of these isolated sites compared with those on the continental shelf which have only been established in the last 15 000 yr and still retain differences arising from the initial founder events. Received: 15 January 1997 / Accepted: 15 August 1997     

Genetic structure of local populations and divergence between growth forms in a clonal invertebrate,the Caribbean octocoral Briareum asbestinum

Although the genetic structure of many populations of marine organisms show little deviation from panmixia, in those marine species with limited larval dispersal, patterns of microgeographic genetic differentiation may be common. The octocoral Briareum asbestinum should show local population differentiation because colonies reproduce asexually by fragmentation, most matings occur between colonies in very close proximity, and the sexually produced larvae and sperm appear to disperse only short distances. Variability in secondary chemistry of individual B. asbestinum colonies from different populations in close proximity also suggests local population differentiation. We determined the genetic composition of local populations by surveying allozyme variation of three shallow and two deep populations within a 300 m² area at San Salvador Island, Bahamas and at a site 161 km away on Little San Salvador, Bahamas in July 1990. As B. asbestinum occurs as either an erect branching form or an encrusting mat often at the same sites, we sampled both morphs to examine the extent of genetic exchange between them. Five of 21 loci were polymorphic and most populations showed a deficit of heterozygotes. Allele frequencies differed significantly between morphs at each site where they occurred together. The mean genetic distance (D=0.065) between morphs is consistent with the interpretation that the two morphs are genetically isolated. Despite the close spatial proximity of the San Salvador populations, both the branching and encrusting morphs showed significant genetic heterogeneity among neighboring populations. Similarly, pooled allelic frequencies for samples collected from the islands of San Salvador and Little San Salvador differed significantly at 1 locus for the branching morph and at 3 out of 5 loci for the encrusting morph.     

Interaction of alongshore sediment transport and habitat conditions at Laguna La Mancha, Veracruz, Mexico

The habitats of La Mancha Lagoon, located midway along the coast of Veracruz, Mexico, are responding to the change of sediment supply reaching its primary inlet at the Gulf of Mexico. Until several decades ago, an abundant alongshore supply of sediment created a periodic opening and closing of the La Mancha inlet. The hydrologic regime of the lagoon consisted of raised water level and lower salinity during the closures, whereas the open inlet favored lower water level, higher salinity, and sediment accumulation in the flood tidal delta. Currently, diminished alongshore sediment supply has affected the inlet morphology and the discharge regime. Associated with the reduced sediment supply, the inlet is open longer in its periodic cycle, the water level variation is reduced, the salinity contrasts are reduced, and the rate of sedimentation in the flood-tide delta is increased. This combination of alterations to the inlet area is changing the flooding regime and affecting the conditions in a very well-developed mangrove habitat at the lagoon margins as well as conditions within the aqueous portions of the lagoon. Management options produce a conflict between supporting the direction of change or preserving the existing habitats.     

Genetic structure in lagoons: the effects of habitat discontinuity and low dispersal ability on populations of <Emphasis Type="Italic">Atherina boyeri</Emphasis>

Atherina boyeri is a common inhabitant of the Mediterranean brackish and coastal waters. Several recent studies proposed the recognition of three distinct forms/species (one lagoon/freshwater and two marine) within an A. boyeri complex. In this study, we used 10 newly available microsatellite loci to better define the genetic structure of the lagoon form. A total of 428 fishes from 11 lagoons along the Italian and Albanian coasts and three Italian lakes were analyzed. A pronounced genetic spatial structure was confirmed for all considered geographic scales (near, medium and broad). A restricted gene flow was detected, suggesting reduced population connectivity. The comparison of present data and previous studies, based on mtDNA and RAPD markers, allowed a preliminary identification of the roles of recent factors (gene flow, habitat discontinuity and water currents) and past events (e.g., Pleistocene glaciations) in shaping the pattern of genetic differentiation.     

Symbiont diversity within the widespread scleractinian coral<Emphasis Type="Italic"> Plesiastrea versipora</Emphasis>, across the northwestern Pacific

The molecular diversity of symbiotic dinoflagellates associated with the widespread western Pacific coral Plesiastrea versipora was explored in order to examine if associations between reef-building corals and symbiotic dinoflagellates change with environment. Several ribosomal DNA genes with different evolutionary rates were used, including the large subunit (28S), the 5.8S region and the internal transcribed spacers (ITS). The phylogenetic analysis of the 28S and 5.8S rDNA regions indicated that a single endosymbiont species, highly related to one of the species of Symbiodinium in clade C (= Symbiodinium goreaui, Trench et Blank), associates with P. versipora along the Ryukyu Archipelago. The persistence of the same endosymbiont within P. versipora across this wide array of latitudes may be a result of such features as the Kuroshio Current, which brings tropical temperatures as far north as Honshu, Japan. Analysis of the faster evolving ITS rDNA region revealed significant genetic variability within endosymbionts from different populations. This variation was due to a high degree of interpopulation variability, based on the proportion of pairwise variation detected among the populations (0.95% approximately). By comparison with other studies, the results also indicate that some ITS1 haplotypes from P. versipora endosymbionts seem to be widely distributed within the western Pacific Ocean, ranging from the Great Barrier Reef to the northeast of the China Sea.     

The stochastic demography of two coexisting male morphs

Abstract. If genetically distinct morphs coexist under a range of natural conditions, they should have equal long-run fitnesses across a wide range of different stochastic environments. In other words, the sequence and frequency of good and bad environments should not substantially impact long-run growth rates. When different morphs have contrasting life histories that vary with environmental conditions, however, it seems improbable that growth rates can be equivalent across a range of stochastic environments without invoking a strong stabilizing mechanism to explain their persistence. As yet, there has been no research characterizing the long-run stochastic growth rate (lambdaS) of different morphs across a wide range of stochastic environments. Assuming density independence, we show that the two genetic male morphs in the bulb mite (Rhizoglyphus robini-fighters, which are able to kill other mites, and benign scramblers-have similar lambdas in different Markovian environments (different simulated random sequences of good and bad habitats). Elasticity analyses revealed that Xs was most sensitive to perturbation of adult survival rate. A slight (biologically and statistically realistic) increase in scrambler adult survival equalized scrambler and fighter X,. The fitness equivalence of the two morphs suggests that stabilizing mechanisms, such as density or frequency dependence, required to maintain their coexistence, are weak. We advocate that stochastic demography can offer a powerful approach to identify and understand the circumstances under which genetic polymorphisms can be maintained in stochastic environments.     

Conservation genetics of <Emphasis Type="Italic">Lumnitzera littorea</Emphasis> (Combretaceae), an endangered mangrove,from the Indo-West Pacific

Mangrove forests, with their ecological significance and economic benefits, are vital inter-tidal wetland ecosystems. Lumnitzera littorea (Combreataceae) is a non-viviparous mangrove distributed in tropical Asia and North Australia. Due to natural and human impacts, populations of this species have been isolated, fragmented, and highly disturbed. In China, L. littorea is an endangered species, restricted to small regions of Hainan Island. The genetic composition of five populations of this species from the Indo-West Pacific (South China, Malay Peninsula, Sri Lanka, North Australia) was assessed using inter simple sequence repeat (ISSR) makers. At the species level, expected mean heterozygosity (He) was 0.240 with 75.6% of loci polymorphic (P). However, genetic variation was much lower at the population level (P = 37.1%, He = 0.118). A high coefficient of gene differentiation (Gst = 0.515) and low level of gene flow (Nm = 0.470) indicated significant genetic differentiation among populations. AMOVA also indicated that more than half the total variation (58.4%) was partitioned among populations. The high degree of differentiation observed among populations emphasizes the need for appropriate conservation measures that incorporate additional populations into protected areas, and achieve the restoration of separate, degraded populations.     

Lower genetic diversity in the limpet <Emphasis Type="Italic">Patella caerulea</Emphasis> on urban coastal structures compared to natural rocky habitats

Human-made structures are increasingly found in marine coastal habitats. The aim of the present study was to explore whether urban coastal structures can affect the genetic variation of hard-bottom species. We conducted a population genetic analysis on the limpet Patella caerulea sampled in both natural and artificial habitats along the Adriatic coast. Five microsatellite loci were used to test for differences in genetic diversity and structure among samples. Three microsatellite loci showed strong Hardy–Weinberg disequilibrium likely linked with the presence of null alleles. Genetic diversity was significantly higher in natural habitat than in artificial habitat. A weak but significant differentiation over all limpet samples was observed, but not related to the type of habitat. While the exact causes of the differences in genetic diversity deserve further investigation, these results clearly point that the expansion of urban structures can lead to genetic diversity loss at regional scales.     

Multiple trans-Arctic passages in the red alga Phycodrys rubens: evidence from nuclear rDNA ITS sequences

In order to investigate how episodes of geological and climatic change have influenced the distribution and evolutionary diversification of Arctic to cold temperate-North Atlantic seaweed species, intraspecific genetic variation was analyzed among isolates of the sublittoral, benthic red alga Phycodrys rubens (collected between June 1992 and January 1994). Rooted phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS) sequences and the plastid encoded Rubisco spacer sequences suggest that P. rubens invaded the North Atlantic from the Pacific shortly after the opening of the Bering Strait (3 to 3.5 million years ago), colonizing both the western and eastern Atlantic coasts. Based on these data we further hypothesize that P. rubens survived along the European coasts during the more recent Pleistocene glaciations, while becoming locally extinct along the North American Atlantic coasts. Following retraction of the last ice sheet, the western Atlantic coast was colonized a second time from the Pacific. The presence of two distinct genetic types (based on ITS and Rubisco sequences) along the European coasts is postulated to be a result of isolation and subsequent differentiation. This is likely because ice-free areas are known to have existed in northern Scotland and Norway during the last glaciation. The presence of an East Atlantic genetic type along the West Atlantic coast is believed to be a recent introduction (caused by human activity) of P. rubens to Newfoundland.     

Habitat-specific variation and performance trade-offs in shell armature of New Zealand mudsnails

Studies documenting phenotypic variation among populations show that ecological performance in one activity is sometimes traded off against another. Identifying environment-specific costs and benefits associated with performance trade-offs is fundamental to knowing how conflicting selection pressures shape phenotype-environment matching in populations. We studied phenotypic variation in shell armature (spininess) of the New Zealand mudsnail, Potamopyrgus antipodarum (Gray), and explored how this variability relates to performance trade-offs between flow resistance and predator deterrence. Smooth- and spiny-shell morphotypes exist in populations in New Zealand streams and lakes, but the patterns and correlates of spatial variation of these phenotypes, and the possible hydrodynamical constraints and antipredatory benefits associated with spiny shell armature, are unknown. Samples from 11 rivers and nine lakes on the South Island showed that, on average, nearly 70% of snails in streams were smooth-shelled, whereas >80% of snails in lakes were spiny, suggesting dissimilar selective pressures between habitats. A laboratory flume experiment revealed that spines collected seston (i.e., suspended algae) at current speeds <40 cm/s, making spiny morphs more prone to flow-induced dislodgment than smooth morphs. However, a fish feeding experiment showed that one benefit of spines on shells was a decrease in predation risk from the common bully (Gobiomorphus cotidianus), a widespread predator of mudsnails in both streams and lakes. All snails egested by bullies were dead, further suggesting that these fishes may exert strong lethal effects on mudsnail populations in nature. Spine expression in lakes also appeared to be temperature related. We conclude that functional trade-offs between risk of flow-induced dislodgment and risk of fish predation affect shell armature frequencies of Potamopyrgus in freshwater habitats.     

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.