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.     

Morphological divergence of Eastern Pacific and Caribbean isopods: effects of a land barrier and the Panama Canal

The Isthmus of Panama rose approximately 3 million years before the present (mybp) and isolated biotas in the tropical eastern Pacific from those in the Caribbean Sea. Populations that were split by the Isthmus and have evolved in allopatry since that time are known as geminates. The surf zone/beach isopod Excirolana braziliensis Richardson was examined between 1984 and 1989 to test the hypothesis that divergence in geminate isopod morphology has occurred, and that geminate divergence is greater than divergence between local populations from the same coastline. Three morphs of Excirolana braziliensis, one in the Caribbean and two in the eastern Pacific, were discovered using numerical taxonomic methods that adjust for body size. The two Pacific morphs have overlapping large-scale distributions, but those morphs are segregated on a smaller scale by beach. We inferred that one Pacific morph and one Caribbean morph were geminates, based on their relative similarity in shape, their geographical ranges, and natural history information about the organism's dispersal capabilities. The origin of the third morph probably predates the Isthmus of Panama, given its relative dissimilarity from the geminate morphs. The presumed geminates differ primarily with respect to the rostrum, antennae and one male reproductive structure. Divergence between geminates is greater than divergence between local populations of any morph along a coastline. Because only one morph occurs in the Caribbean, that region contains less morphological variation than the eastern Pacific, which contains two morphs. There was weak evidence that some introductions may have taken place in the last century from the Caribbean to the Pacific; however, introductions have not masked the pattern of divergence that has developed over millions of years.     

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     

Social,environmental and genetic factors in the ontogeny of phenotypic differentiation in a lizard with alternative male reproductive strategies

Summary Adult male tree lizards, Urosaurus ornatus, practise alternative (territorial or sneaker/satellite) reproductive strategies that are correlated with differences in throat color and body size. In this study we raised tree lizards from hatching in the laboratory to examine the question of whether the phenotypic expression of secondary sex coloration and body size can be facultatively influenced by social or abiotic environmental factors. We compared males reared in the laboratory under different social and environmental conditions to males in the field and found no effect of different conditions on phenotypic differentiation (Figs. 2–4). Thus, phenotypic differences between morphs probably result largely from nonfacultative expression of different genotypes. This suggests that alternative male morphs practise a mixed evolutionary stable strategy (ESS) rather than one morph making the best of a bad situation. However, in the context of ESS theory it is difficult to explain our further result that the nonterritorial morph in this species grows faster and reaches a larger adult body size than the territorial morph (Fig. 5).     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

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.     

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.     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Distribution pattern, reproductive traits, and molecular analysis of two coexisting vermetid gastropods of the genus Petaloconchus: a Caribbean endemic and a potential invasive species

Vermetid gastropods are characterized by complex taxonomy and unusual ecology. A survey of the fouling community in the intake channel at the Planta Centro Power Plan in Puerto Cabello, Venezuela, found massive colonies of vermetid gastropods of the genus Petaloconchus. We arbitrarily named two prevalent varieties as black and brown-orange morphs, distinguishing based on the color of their soft bodies. Spatial distribution was different for the morphs. The black morph was present along the jetty, with higher average densities in the shallower intertidal area (410 vs. 143 ind m⁻²), while the brown-orange morph was only present at the initial part of the channel (μ = 83 ind m⁻²). Both produced small eggs (142 vs. 180 μm diameter on average), with the orange-brown eggs being slightly larger, but the morphs differed in other reproductive aspects. The brown-orange morph produced significantly fewer capsules (up to nine simultaneously per female) with fewer embryos (average of 27 viable embryos/capsule) that hatched at a larger size (577 μm on average), allocating about 17.64% of the initial egg production to nurse eggs. In contrast, the black morph produced up to 14 capsules simultaneously, hatched an average of 178 veligers of 212 μm, and produced no nurse eggs. The intra-capsular development reached a more advanced stage in the brown-orange morph than in the black one. Several cytochrome subunit I (COI) and 16S rRNA (16S) haplotypes were found for the black morph, when compared to only one haplotype for both genes present for the brown-orange morph. Preliminary phylogenetic analyses separated the morphs into different clades, supported by robust bootstrap values and posterior probabilities (>98). Our results indicate that the morphs are two different species: the black morph was identified as Petaloconchus cf. varians and the brown-orange morph as a non-described species, Petaloconchus sp. (orange). The first is endemic to the Caribbean, while the second is potentially an introduced species.     

Intermorph breeding and the potential for reproductive isolation in polymorphic mole salamanders (Ambystoma talpoideum)

The study of reproductive isolation as a prerequisite to sympatric speciation has been limited by the focus on species that have already experienced such isolation. However, a complete understanding of how such processes evolve depends on observing taxa before they complete the speciation process. We studied the potential for sexual isolation in the polyphenic mole salamander, Ambystoma talpoideum, using a series of laboratory and field experiments. This species consists of aquatic paedomorphic adults and terrestrial metamorphic adults which are exhibited by both sexes and which mate in the same aquatic habitat. Previous field studies on this species suggested that intermorph breeding would be less common during the winter months, because paedomorphic adults begin breeding in early autumn and thus may have less energy available for reproduction in the winter. Laboratory experiments conducted during the winter showed that the mating behavior of paedomorphic males occurred at a much lower frequency than that of metamorphic males. In contrast, field experiments that best mimicked natural conditions revealed symmetric intermorph breeding and included multiple paternity shared among males of each morph. This and other studies suggest that there is little evidence of sexual isolation among morphs based on behavioral interactions alone. However, the potential for partial isolation still occurs because of temporal and spatial differences in the frequencies of each morph in nature. Our results suggest that further studies on this system, and other similar polyphenisms, may provide valuable insight into the mechanisms that underlie the evolution of reproductive isolation.     

Lack of innate preference for morph and species identity in mate-searching <Emphasis Type="Italic">Enallagma</Emphasis> damselflies

Insect mate recognition is often viewed as stereotypic, innate, and species-specific. However, male damselflies can learn to identify female-specific color morphs as potential mates. A suite of male mimicry hypotheses assume that heteromorphic females, which differ from males in color pattern, are more easily recognized as “female” and thus lack the inherent, anti-harassment advantage that the more male-like signal provides for andromorphs. Using two measures of male preference, we investigated whether naïve males have a preexisting sensory bias for a given morph color in Enallagma civile, a species that appeared to exhibit extreme plasticity in morph expression across generations within a breeding season. E. civile males raised in the absence of females exhibited no preference for either morph, whereas males raised with one female type exhibited a learned sensory bias for that morph. Male Enallagma also lacked a bias toward conspecific females over a congeneric sister species. In a naturally naïve population of Enallagma ebrium, males reacted sexually to both morphs of Enallagma hageni as often as they did to conspecific females, whose thoracic spectra were nearly identical with those of E. hageni. Moreover, despite the similar thoracic spectra of males and andromorphs, both of which reflected UV, males rarely reacted sexually to other males. Our results falsified implicit assumptions of male mimicry hypotheses, supported learned mate recognition, and suggested a scenario for speciation via sexual conflict.     

Dimorphism and possible sex change in copepods of the family Calanidae

Adult females of 14 of 25 species of the family Calanidae were found to be dimorphic with respect to the number of aesthetases on the first antenna. The trithek morph, in which most antennal segments bear a complement of one aesthetasc and two setae, appears to be the typical female phenotype. The quadrithek morph is less common and, as in males, odd-numbered segments 2b, 3, 5, 7 and 9 carry quadritheks, i.e., two aesthetascs as well as two setae. Segment 21 in the quadrithek female bears an aesthetasc that is absent in the trithek female. Male antennal segments are distinguished from those of trithek and quadrithek females by pronounced morphological differences in aesthetasc shape and size as well as fusion of one or more pairs of antennal segments. The quadrithek morph usually comprised only a small proportion (<10%) of a local population. Quadrithek morphs were found in tropical and subtropical genera (Cosmocalanus and Nannocalanus), broadly ranging genera (Calanoides), as well as in cool-water lineages (Calanus s.s., Calanus s.l. cristatus, C. s.l. plumchrus, C. s.l. propinquus and C. s.l. tonsus). Maximum frequencies of quadrithek morphs (10 to 12%) in Calanus pacificus californicus occurred during the upwelling season from late winter into spring. No indication of sexual or antennal dimorphism was found in CV C. pacificus californicus sexed by examination of gonad and gonaduct. The quadrithek dimorph appears to be the product of sex change by the larger-sized late-juvenile potential male. The evolutionary and ecological significance of this presumptive hypothesis is that the larger-sized potential male reaching adulthood one or more weeks before maturation of genotypic females may increase its reproductive output by sex change in the course of the final molt to mature as a functional female. Given the sex-change hypothesis, morphogenesis of secondary sexual structures would appear to be controlled by androgenic-like secretions from the genital tract, as has been shown in malacostrocans.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.     

Nemertean,copepod, and amphipod symbionts of the dimorphic ascidian Pyura stolonifera near Melbourne,Australia: specificities to host morphs,and factors affecting prevalences

On the central coast of Victoria. Australia, the dimorphic ascidian Pyura stolonifera (Heller, 1878) harbors three endosymbionts: the nemertean Gononemertes australiensis Gibson, 1974, the copepod Doropygus pulex (Thorell, 1859), and the amphipod Paraleucothoe novaehollandiae (Haswell, 1880). The specificities of these symbionts to two host colour morphs were studied during 1989 to 1991 as part of a multidisciplinary investigation aimed at determining whether the two morphs are genetically distinct. Distributional surveys revealed that nemerteans and copepods occur only in yellow and brown ascidians, respectively, and that amphipods live in both forms. These specificities held true not only when the two morphs were in allopatry, but also in sympatry. These observations, especially the sympatric data, suggest that the two host morphs might be genetically distinct. For example, the two morphs might have different genetically encoded internal milieus that favour the survival of nemerteans in yellow ascidians, and copepods in brown hosts. In transplant experiments, which involved moving ascidian morphs within and between habitats, the wrong symbionts never colonised the wrong hosts. These results, although consistent with the hypothesis of genetic maintenance of specificity, were deemed inconclusive because of the difficulty of establishing reliable controls (i.e. vacant hosts). The relationships between symbiont prevalences and several factors (season, year, site within host, host individual, host habitat, host size/age, host breeding condition, and co-occurrence of other symbiont species) were also analysed. Both simple (e.g. greater prevalences for large hosts) and complex (e.g. prevalence x season x gonad state of host) interactions were detected for all three symbiont species. These are among the very few quantitative analyses of factors affecting prevalences of ascidicolous nemerteans and amphipods. The present report identifies one of very few definite nemertean-ascidian symbioses. Since no differences in gross condition were ever noticed between occupied and vacant hosts, it is suggested that all three symbionts are commensals rather than parasites or mutuals.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

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.     

Acanthochromis polyacanthus,a fish lacking larval dispersal,has genetically differentiated populations at local and regional scales on the Great Barrier Reef

Acanthochromis Gill is a monotypic genus within the damselfish family Pomacentridae, erected for an unusual species [A. polyacanthus (Bleeker)] that uniquely lacks larval dispersal. Instead, offspring are reared in the parental territory, in the manner of cichlids, and fledged into the surrounding habitat. Phenotypic and genotypic variation was surveyed on the basis of body colouration and 7 polymorphic loci in 19 populations from 5 regions of the central and southern Great Barrier Reef (GBR). Variation in both characters was found at regional and local scales. Two colour morphs were recognised: a bicoloured morph from the three northern regions and a uniform dark morph from the two southern regions. Isozyme analysis showed a similar pattern with greatest variation between the different morphs, but also with significant variation at both regional and local scales within morphotypes. Heterozygosity was maximal in the central populations, which, together with other measures of variability, suggests a mixing of separate gene pools in this region and denies species status to the two morphotypes despite numerous fixed differences in allele frequencies between the most distant populations. The presence of fixed differences in multiple alleles between populations separated by 1000 km indicates negligible gene flow over such distances and long isolation of these gene pools. These patterns may reflect recolonisation of the GBR after the last sea-level rise by fish from two stocks. Founder effects and random drift in small populations after colonisation are probably the major sources of the local and regional variations observed at smaller spatial scales. This diversity has been maintained among populations at all scales by the very low levels of gene flow possible without an effective strategy for larval dispersal between coral reefs.     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Temporal reproductive isolation and gametic compatibility are evolutionary mechanisms in the<Emphasis Type="Italic"> Acropora humilis</Emphasis> species group (Cnidaria; Scleractinia)

Patterns of interbreeding between individuals are fundamental to the structure and maintenance of evolutionary boundaries between species. In corals, both hybridisation and reproductive isolation appear to be important evolutionary mechanisms. In this study, I examine evolutionary boundaries using morphological, molecular and reproductive criteria within the Acropora humilis species group at Lizard Island on the Great Barrier Reef, Australia. Five species and seven morphs are recognised on the basis of morphological appearance of features traditionally used to identify corals of the genus Acropora. In a molecular phylogenetic analysis, I examine relationships for the mitochondrial DNAs putative control region, using maximum-parsimony and maximum-likelihood methods. The reproductive criteria explore whether species and morphs are reproductively isolated on the basis of temporal or fertilisation barriers. Timing of gamete maturity is surveyed for each species and morph, from the month prior to and 3 months after the mass spawning. Time of spawning is documented at the levels of night and hour of spawning, and time taken for egg-sperm bundles to separate. Laboratory fertilisation experiments tested the potential of species and morphs to interbreed. High levels of intraspecific and extremely low or zero fertilisation levels between the five species indicated that they are valid species. Based on the combined assessment of morphological, molecular and reproductive criteria, A. humilis and A. gemmifera appear to be the most closely related species, which are most closely related to the remaining species in the following order: A. samoensis, A. monticulosa and A. digitifera. Evidence derived from one or more of these criteria suggest that the morphs (1) are at various stages of divergence from the species with which they share morphological characters, and (2) may indicate possible zones of speciation and hybridisation. Identification of morphs avoided the possibility of taxonomic error and was essential for accurate interpretation of evolutionary boundaries. Confirmation of morphology as an informative character of evolutionary boundaries is of great significance because most coral research projects rely on morphology as the primary tool for identification of species.Communicated by M.S. Johnson, Crawley