Contrast Between Solitary and Clonal Lifestyles in the Sea Anemone Anthopleura elegantissima

The common intertidal sea anemone Anthopleura elegantissima(Brandt) occurs in two forms the clonal aggregating form andthe solitary form The obvious differences between the two apparentlyresult from the presence of asexual reproduction in the clonalaggregating form and its absence in the solitary form resultingdifferences in growth form suit the two anemones to differentlifestyles The clonal form is well suited to life higher inthe mid intertidal Asexual reproduction resulting in moderatelysmall individual size and close association with clonematesimproves its resistance to physical stress (drag and desiccation)and makes it a superior competitor in exposed habitats higherin the intertidal where species diversity and predator pressureare low The larger solitary animals live in more protected microhabitatslower in the intertidal and subtidal and are probably more resistantto predation and less able to withstand physical stress or intenseintraspecific competition The two forms also have differentbiogeographic ranges While the clonal aggregations are quitecommon at rocky sites at least as far north as Vancouver Islandin British Columbia populations of solitaries have not beenfound north of Point Reyes California Apparently the two formsare re productively isolated phenotype frequencies are verydifferent for clonals and solitaries living at the same locationswhile the between-sample variation within each form is relativelysmall These differences in phenotype frequencies biogeographicrange and microhabitat suggest that the clonal anemones andthe solitary anemones known as Anthopleura elegantissima areactually a sibling species pair.     

Growth, reproduction and survival of a tropical sea anemone (Actiniaria): benefits of hosting anemonefish

The ecological performance of the sea anemone Heteractis magnifica was examined during a 36-month experiment with respect to season and the presence and numbers of a mutualist (orange-fin anemonefish Amphiprion chrysopterus). Anemones primarily grew during the autumn, with most asexual reproduction occurring in winter; mortality was not strongly seasonal. Individual growth rates did not differ between anemones harboring one or two anemonefish, but these rates were three times faster than for anemones lacking Amphiprion. Anemones with two anemonefish had the highest fission rate, whereas those without anemonefish had the lowest. By contrast, anemones that were not defended by anemonefish suffered higher-than-expected mortality. As a consequence, anemones with two Amphiprion had the greatest net increase in surface area, and those lacking anemonefish had a negligible gain that was statistically indistinguishable from zero after three years. Anemonefish not only enhanced anemone survivorship as previously believed, they also fostered faster growth and more frequent asexual reproduction.     

Geographic differences in vertical connectivity in the Caribbean coral Montastraea cavernosa despite high levels of horizontal connectivity at shallow depths

The deep reef refugia hypothesis proposes that deep reefs can act as local recruitment sources for shallow reefs following disturbance. To test this hypothesis, nine polymorphic DNA microsatellite loci were developed and used to assess vertical connectivity in 583 coral colonies of the Caribbean depth‐generalist coral Montastraea cavernosa. Samples were collected from three depth zones (≤10, 15–20 and ≥25 m) at sites in Florida (within the Upper Keys, Lower Keys and Dry Tortugas), Bermuda, and the U.S. Virgin Islands. Migration rates were estimated to determine the probability of coral larval migration from shallow to deep and from deep to shallow. Finally, algal symbiont (Symbiodinium spp.) diversity and distribution were assessed in a subset of corals to test whether symbiont depth zonation might indicate limited vertical connectivity. Overall, analyses revealed significant genetic differentiation by depth in Florida, but not in Bermuda or the U.S. Virgin Islands, despite high levels of horizontal connectivity between these geographic locations at shallow depths. Within Florida, greater vertical connectivity was observed in the Dry Tortugas compared to the Lower or Upper Keys. However, at all sites, and regardless of the extent of vertical connectivity, migration occurred asymmetrically, with greater likelihood of migration from shallow to intermediate/deep habitats. Finally, most colonies hosted a single Symbiodinium type (C3), ruling out symbiont depth zonation of the dominant symbiont type as a structuring factor. Together, these findings suggest that the potential for shallow reefs to recover from deep‐water refugia in M. cavernosa is location‐specific, varying among and within geographic locations likely as a consequence of local hydrology.     

The annual gametogenic cycle of the sea anemone Metridium senile from the Gulf of Maine

A year-long study was conducted to quantify the reproductive cycle of the clonal sea anemone, Metridium senile, an important member of the benthic community in the Gulf of Maine (GOM). M.senile is an abundant sessile invertebrate that forms identifiable clonal aggregations that are maintained by pedal laceration but individuals can also reproduce sexually. Specimens were collected monthly from a shallow (< 10 m) subtidal site and the reproductive cycle of male and female anemones was described on a seasonal basis using histological preparation of the gonads and light microscopy. Gametogenesis was determined for both sexes and showed that females undergo spawning during the seasonal transition from summer to fall, while males have mature sperm within testicular cysts in the winter, spring, and summer. Environmental data recorded at the study site shows that the spawning period occurs during a period of peak water temperature (August-September), and rapid gametogenesis occurs during periods of high food availability (April). The low percentage of reproductively mature anemones and the dominance of asexual reproduction in this population are believed to be the result of the high flow regime at this site.     

POPULATION STRUCTURE OF A CLONAL GORGONIAN CORAL: THE INTERPLAY BETWEEN CLONAL REPRODUCTION AND DISTURBANCE

Clonality is a common feature of plants and benthic marine organisms. In some cases clonal propagation results in a modest increase in population density, while in other cases dense populations may be generated by the propagation of only a few clones. We analyzed the population structure of the clonal gorgonian Plexaura kuna across several reef habitats with a range of disturbance regimes in the San Blas Islands, Panama, and the Florida Keys, U.S.A. Using multilocus DNA fingerprinting to distinguish clones, we estimated that clones ranged in size from single individuals to 500 colonies. The number of genotypes identified on nine reefs ranged from three to 25. Population density and clonal structure varied markedly among reefs with G_O:G_E ranging from 0.03 to 1.00. On some reefs vegetative reproduction transformed P. kuna from a rare species to the numerically most abundant gorgonian. The effect of clonal propagation on P. kuna population structure was dependent on interactions between fragmentation and the reef environment (disturbance regime, substratum). We present a generalized model relating population structure of clonal species to disturbance and the mode of vegetative propagation. Disturbance promotes colony propagation and skews the size-frequency distribution of clones among P. kuna and many species that propagate via fragmentation. Propagation of these species is promoted by disturbance (disturbance sensitive), and they tend to have clones that are dispersed across local sites. Species that fragment and have dispersed clones, have high genotypic diversity in habitats with low levels of disturbance. Genotypic diversity then decreases at intermediate disturbance and increases again at the highest disturbance levels. Clonal species that do not rely on disturbance for vegetative propagation (disturbance insensitive) generally do not disperse and form aggregated clones. Among these taxa disturbance has a greater affect on individual survival than on propagation. Genotypic diversity is directly related to the level of disturbance until very high levels of disturbance, at which time genotypic diversity declines.     

Internal brooding of clonal propagules by a sea anemone, Anthopleura sp.

Abstract. We investigated the seasonal prevalence of reproductive activities and of the development of brooded propagules in an intertidal sea anemone, Anthopleura sp., on the rocky shore of Mutsu Bay, in northern Japan. A monthly examination of anemones, by dissection and histological techniques, revealed no sign of gonad development, but did reveal that they produce and internally brood propagules throughout the year. Release of propagules was observed in the field. This anemone population appears to be entirely asexual and agametic, and may persist solely through clonal propagation.     

Polyploidy and microsatellite variation in the relict tree Prunus lusitanica L.: how effective are refugia in preserving genotypic diversity of clonal taxa?

Refugia are expected to preserve genetic variation of relict taxa, especially in polyploids, because high gene dosages could prevent genetic erosion in small isolated populations. However, other attributes linked to polyploidy, such as asexual reproduction, may strongly limit the levels of genetic variability in relict populations. Here, ploidy levels and patterns of genetic variation at nuclear microsatellite loci were analysed in Prunus lusitanica, a polyploid species with clonal reproduction that is considered a paradigmatic example of a Tertiary relict. Sampling in this study considered a total of 20 populations of three subspecies: mainland lusitanica (Iberian Peninsula and Morocco), and island azorica (Azores) and hixa (Canary Islands and Madeira). Flow cytometry results supported an octoploid genome for lusitanica and hixa, whereas a 16‐ploid level was inferred for azorica. Fixed heterozygosity of a few allele variants at most microsatellite loci resulted in levels of allelic diversity much lower than those expected for a high‐order polyploid. Islands as a whole did not contain higher levels of genetic variation (allelic or genotypic) than mainland refuges, but island populations displayed more private alleles and higher genotypic diversity in old volcanic areas. Patterns of microsatellite variation were compatible with the occurrence of clonal individuals in all but two island populations, and the incidence of clonality within populations negatively correlated with the estimated timing of colonization. Our results also suggest that gene flow has been very rare among populations, and thus population growth following founder events was apparently mediated by clonality rather than seed recruitment, especially in mainland areas. This study extends to clonal taxa the idea of oceanic islands as important refugia for biodiversity, since the conditions for generation and maintenance of clonal diversity (i.e. occasional events of sexual reproduction, mutation and/or seed immigration) appear to have been more frequent in these enclaves than in mainland areas.     

Population Structures in Rhododendron metternichii var. hondoense Assessed with Microsatellites and their Implication for Conservation

Rhododendron metternichii var. hondoense is its great morphological variation. Individuals may have only one erect stem or may have multiple creeping stems, implying that some of them recruit vegetatively. Aims of this study are to ascertain whether a population of R. metternichii var. hondoense consists of clonal plants, and to evaluate relative importance of sexual and asexual recruitments in regard to its conservation. Six microsatellite loci were analyzed in two populations growing in different habitats. One was in a mesic valley consisting of many sprouting and creeping individuals with few seedlings, and the other on a mountain ridge consisting of single stem individuals with many seedlings. Sufficient polymorphisms were found to be present even in the mesic valley population that consisted of many sprouting and creeping stems, indicating that in the past the population was maintained by both sexual and asexual reproduction. The scarcity of seedlings at the mesic valley was due to dense litter cover and low bryophyte mat cover which may be caused by changes in traditional management systems. Required conservation measures are discussed based on these results. Received 8 July 1999/ Accepted in revised form 26 August 1999     

Exploring the role of asexual multiplication in poplar rust epidemics: impact on diversity and genetic structure

Fungal plant pathogens, especially rust fungi (Pucciniales), are well known for their complex life cycles, which include phases of sexual and asexual reproduction. The effect of asexual multiplication on population genetic diversity has been investigated in the poplar rust fungus Melampsora larici‐populina using a nested hierarchical sampling scheme. Four hierarchical levels were considered: leaf, twig, tree and site. Both cultivated and wild poplar stands were sampled at two time points at the start and end of rust epidemics. A total of 641 fungal isolates was analysed using nine microsatellite markers. This study revealed that the genetic signature of asexual multiplication in the wild poplar stand was seen only at lower hierarchical levels (leaf and twig). Moreover, we observed an erosion of clonal structure through time, with an increase in both gene and genotypic diversity. New genotypes contributed to host infection over time, which demonstrates the importance of allo‐infection in the epidemic process in this host‐pathogen system. Compared with the wild stands, the nearly lack of detection of clonal structure in the cultivated stands reflects the higher infection level on cultivated poplars. More generally, this genetic analysis illustrates the utility of population genetics approach for elucidating the proportion of asexual reproduction in the multiplication of isolates during an epidemic, and for proper quantification of asexual dispersal in plant pathogens.     

Molecular discrimination and phylogeographic patterns of clones of the parthenogenetic gecko Lepidodactylus lugubris in the Japanese Archipelago

Vertebrates usually reproduce sexually in which males and females contribute their offspring genome and produce genetically diverse offspring. However, some of them are asexual without genetic contribution from males. The nocturnal gecko, Lepidodactylus lugubris, is all females and reproduces parthenogenetically. This gecko is known to consist of diploid and triploid clones in the tropical and subtropical regions, which can be identified by their dorsal marking patterns, ploidy, and protein polymorphism. This gecko is also distributed in the southern parts of Japan, and several clones have been reported. In this study, we investigated the origins and genetic diversity of Japanese L. lugubris by clonal discrimination using microsatellite and mitochondrial DNA analyses. A total of 748 individuals were collected from 21 islands of five island groups (Ogasawara, Okinawa, Miyako, Yaeyama and Daito Islands) and 17 clones were distinguished genetically. Mitochondrial cyt b sequences of these clones suggested that they were all closely related and differentiated recently. Clonal diversity was much higher (14 clones) in the Daito Islands than in the other island groups in which only one or two clones coexisted. Judging from the dorsal marking patterns and ploidy known so far, six clones were cosmopolitan and may be colonized from the outside of Japan. However, other 11 clones were endemic to the Daito Islands and explained by possible hybridization between the one female diploid clone and one male diploid clone because other 9 clones were triploid and all had the combinations of polymorphic microsatellite alleles of these female and male diploid clones. Although the males have never been recorded in the Daito Islands, males might appear in the past. These findings contribute to understanding of clonal diversity and dynamics of asexually reproducing animals. If diploid parthenogenetic geckos can produce triploid clones by mating with the diploid males, clonal diversity would increase rapidly in a small island, and such newly produced triploid clones would expand widely.     

Comparative ecology of subtidal actiniarians from the coasts of California and the Gulf of Maine,USA

Shallow subtidal populations of sea anemones occupying hard substrates along the California (Pacific) and Gulf of Maine (Atlantic) coasts of the USA are compared. The diversity of anemones in the southern Gulf of Maine to 35 m is low (4 species) in contrast to similar habitats in California (at least 11 species), but parallel patterns of individual size, distribution and predation impacts occur. Water motion influences depth distribution and size, while predation affects size, population structure and habitat selection. The nudibranch Aeolidia papillosa exerts a similar pressure on population structure in some anemone species in both areas as a size selective predator. The seastar Dermasterias imbricata appears to magnify the selective pressure for large size in the eastern Pacific coastal zone. Interspecific aggression of anemones determines some distribution patterns in areas of overlap.     

Temperate and tropical algal-sea anemone symbioses

Abstract. In this review, we seek to develop new insights about the nature of algal‐sea anemone symbioses by comparing such associations in temperate and tropical seas. Temperate seas undergo pronounced seasonal cycles in irradiance, temperature, and nutrients, while high irradiance, high temperature, and low nutrients are seasonally far less variable in tropical seas. We compare the nature of symbiosis between sea anemones (= actinians) and zooxanthellae (Symbiodinium spp.) in both regions to test tropical paradigms against temperate examples and to identify directions for future research. Although fewer anemone species are symbiotic in temperate regions, they are locally dominant and ecologically important members of the benthic community compared to the tropics. Zooxanthella densities tend to be lower in temperate anemones, but data are limited to a few species in both temperate and tropical seas. Zooxanthella densities are far more stable over time in temperate anemones than in tropical anemones, suggesting that temperate symbioses are more resistant to fluctuations in environmental parameters such as irradiance and temperature. Light‐saturated photosynthetic rates of temperate and tropical zooxanthellae are similar, but temperate anemone hosts receive severely reduced carbon supplies from zooxanthellae during winter months when light is reduced. Symbiont transmission modes and specificity do not show any trends among anemones in tropical vs. temperate seas. Our review indicates the need for the following: (1) Investigations of other temperate and tropical symbiotic anemone species to assess the generality of trends seen in a few “model’ anemones. (2) Attention to the field ecology of temperate and tropical algal‐anemone symbioses, for example, how symbioses function under seasonally variable environmental factors and how zooxanthellae persist at high densities in darkness and winter. The greater stability of zooxanthella populations in temperate hosts may be useful to understanding tropical symbioses in which bleaching (loss of zooxanthellae) is of major concern. (3) Study of the evolutionary history of symbiosis in both temperate and tropical seas. Continued exploration of the phylogenetic relationships between host anemones and zooxanthella strains may show how and why zooxanthellae differ in anemone hosts in both environments.     

Recruitment failure in Florida Keys <Emphasis Type="Italic">Acropora palmata,</Emphasis> a threatened Caribbean coral

Recovery of Acropora palmata from its currently imperiled status depends on recruitment, a process which is poorly documented in existing Caribbean coral population studies. A. palmata is thought to be well adapted to proliferate through the recruitment of fragments resulting from physical disturbances, such as moderate intensity hurricanes. This study monitored fifteen 150 m² fixed study plots on the upper Florida Keys fore-reef for asexual and sexual recruitment from 2004 to 2007. Between July and October 2005, 4 hurricanes passed by the Florida Keys, producing wind speeds on the reef tract of 23 to 33 m s⁻¹. Surveys following the hurricanes documented an average loss of 52% estimated live tissue area within the study plots. The percentage of “branching” colonies in the population decreased from 67% to 42% while “remnant” colonies (isolated patches of tissue on standing skeleton) increased from 11% to 27%. Although some detached branches remained as loose fragments, more than 70% of the 380 fragments observed in the study plots were dead or rapidly losing tissue 3 weeks after Hurricane Dennis. Over the course of the study, only 27 fragments became attached to the substrate to form successful asexual recruits. Meanwhile, of the 18 new, small encrusting colonies that were observed in the study, only 2 were not attributable to asexual origin (i.e., remnant tissue from colonies or fragments previously observed) and are therefore possible sexual recruits. In summary, the 2005 hurricane season resulted in substantial loss of A. palmata from the upper Florida Keys fore-reef from a combination of physical removal and subsequent disease-like tissue mortality, and yielded few recruits of either sexual or asexual origin. Furthermore, the asexual and sexual fecundity of the remaining population is compromised for the near future due to the lack of branches (i.e., “asexual fecundity”) and overall loss of live tissue.     

Genetic variation in sexual and clonal lineages of a freshwater snail

Sexual reproduction within natural populations of most plants and animals continues to remain an enigma in evolutionary biology. That the enigma persists is not for lack of testable hypotheses but rather because of the lack of suitable study systems in which sexual and asexual females coexist. Here we review our studies on one such organism, the freshwater snail Potamopyrgus antipodarum (Gray). We also present new data that bear on hypotheses for the maintenance of sex and its relationship to clonal diversity. We have found that sexual populations of the snail are composed of diploid females and males, while clonal populations are composed of a high diversity of triploid apomictic females. Sexual and asexual individuals coexist in stable frequencies in many ‘mixed’ populations; genetic data indicate that clones from these mixed populations originated from the local population of sexual individuals without interspecific hybridization. Field data show that clonal and sexual snails have completely overlapping life histories, but individual clonal genotypes are less variable than individuals from the sympatric sexual population. Field data also show segregation of clones among depth‐specific habitat zones within a lake, but clonal diversity remains high even within habitats. A new laboratory experiment revealed extensive clonal variation in reproductive rate, a result which suggests that clonal diversity would be low in nature without some form of frequency‐dependent selection. New results from a long‐term field study of a natural, asexual population reveal that clonal diversity remained nearly constant over a 10‐year period. Nonetheless, clonal turnover occurs, and it occurs in a manner that is consistent with parasite‐mediated, frequency‐dependent selection. Reciprocal cross‐infection experiments have further shown that parasites are more infective to sympatric host snails than to allopatric snails, and that they are also more infective to common clones than rare clones within asexual host populations. Hence we suggest that sexual reproduction in these snails may be maintained, at least in part, by locally adapted parasites. Parasite‐mediated selection possibly also contributes to the maintenance of local clonal diversity within habitats, while clonal selection may be responsible for the distribution of clones among habitats. © 2003 The Linnean Society of London. Biological Journal of the Linnean Society 2003, 79 , 165–181.     

Modes of reproduction in sea anemones (Cnidaria,Anthozoa)

The data on different modes of reproduction in sea anemones are generalized. These animals can reproduce sexually in an ordinary way or by parthenogenesis. Asexual reproduction occurs in various forms, such as transverse and longitudinal fission, pedal laceration, or autotomy of tentacles. Specific features of different variants of sexual and asexual reproduction and their combinations in sea anemones from different habitats of the World Ocean are discussed.     

Protection of host anemones by snapping shrimps: a case for symbiotic mutualism?

The sea anemone Bartholomea annulata is an ecologically important member of Caribbean coral reefs which host a variety of symbiotic crustacean associates. Crustacean exosymbionts typically gain protection from predation by dwelling with anemones. Concurrently, some symbionts may provide protection to their host by defending against anemone predators such as the predatory fireworm, Hermodice carunculata, which can severely damage or completely devour prey anemones. Herein we show through both field and laboratory studies that anemones hosting the symbiotic alpheid shrimp Alpheus armatus are significantly less likely to sustain damage by H. carunculata than anemones without this shrimp. Our results suggest that the association between A. armatus and B. annulata, although complex because of the numerous symbionts involved, may be closer to mutualism on the symbiotic continuum.     

The energetics of asexual reproduction: Pedal laceration in the symbiotic sea anemone Aiptasia pulchella (Carlgren, 1943)

Asexual reproduction of the symbiotic sea anemone Aiptasia pulchella (Carlgren, 1943) was studied in the context of energy budgets determined under a variety of light and feeding conditions. Continuous darkness significantly increased rates of pedal laceration (the method of asexual reproduction), while different feeding rates had no significant effect. Digestive assimilation efficiency averaged 63% for whole brine shrimp, and conversion efficiencies (assimilated energy into biomass) averaged 31%. There were no significant differences between the conversion efficiencies of energy obtained from light or food. The index of reproductive effort (RE) for pedal laceration was very low, ranging from 0.004 to 0.044. This low RE associated with a substantial rate of asexual reproduction may, in part, explain the widespread success of certain sea anemones.     

Do reproductive tactics vary with habitat heterogeneity in the intertidal sea anemone Actinia tenebrosa?

Cnidarians display a diverse range of reproductive tactics including sexual and asexual modes of reproduction. Although few studies have looked for intraspecific variation in reproductive tactics, flexible expression of such life-history traits may be favoured in species that occupy a range of habitats. We tested this in the sea anemone Actinia tenebrosa by comparing cycles of reproductive activity and the mode of production of brooded larvae in local populations occupying boulder fields and stable rock platforms. We determined the mode of production of broods from eight rock platforms (separated by up to 1600 km) and two boulder shores on the south east coast of Australia using a combination of allozyme data and four newly characterised microsatellite markers.

We determined seasonal patterns of brooding and gonad development by monthly dissection of 15–30 adults from each of two boulder fields and two stable platforms. Previous genetic studies have shown that populations of A. tenebrosa on rock platforms can be highly clonal, whereas anemones on more heterogeneous boulder habitats display levels of genotypic diversity similar to that expected for sexual reproduction. We genotyped a total of 221 juveniles from 37 brooding adults including 11 broods and 80 juveniles from boulder shores. We did not detect any evidence of sexual production of broods. All brooded juveniles displayed identical multi-locus genotypes to their brood parent irrespective of habitat of origin or location, including 28 broods (200 juveniles) that were heterozygous at one or more locus. Similarly, we found that temporal patterns of gonad formation and brooding were consistent across habitats and locations. We detected 346 mature males, 234 non-reproductive or immature individuals, and no mature females within a total of 580 dissected individuals. These data suggest that the reproductive tactics of A. tenebrosa are essentially fixed and that variation in the genotypic diversity of populations may reflect variation in factors such as the input of sexually derived planktonically dispersed recruits or post-settlement processes. However, the apparent lack of females paradoxically implies that sexual reproduction, and hence recruitment, must be rare or no longer possible within some populations, and highlights the need for long-term studies of these populations.     

Patterns of Juvenile Habitat Use and Seasonality of Settlement by Permit, Trachinotus falcatus

Synopsis Permit, Trachinotus falcatus are economically and ecologically important throughout their range of the Caribbean, subtropical and tropical western Atlantic, and Gulf of Mexico. Despite their economic importance, little is known about the biology and ecology of permit, and most existing information is from Florida. While sufficient information is available to paint a general picture of permit life history, details are lacking for most life stages. For the juvenile life stage, nursery habitats and size and age at settlement have not yet been defined. Although six distinct habitat types (medium energy and low energy windward beaches, leeward beaches, and windward, leeward, and lagoon interior mangrove shorelines) were sampled to determine spatial patterns of habitat use by early juvenile permit at Turneffe Atoll, Belize, Central America, and the Florida Keys, USA, 99% of juvenile permit were found along medium energy windward beaches, indicating their role as nursery habitat for this species. A sub-sample of juvenile permit from Florida was examined to estimate spawning date and age at settlement from otoliths. Size-frequency distributions and otolith age analysis indicate that larval duration is approximately 15 – 20 days, and settlement occurs year-round. Since permit in Florida spawn March through July, from March through September in Cuba, and from February through October in Belize, year-round settlement indicates population connectivity via larval transport. These results lay the foundation for future research on larval supply, population connectivity, and juvenile ecology, and will aid in the ongoing formulation of a conservation plan toward a sustainable fishery for permit.