The evolutionary history of the order Antipatharia (Cnidaria: Anthozoa: Hexacorallia) as inferred from mitochondrial and nuclear DNA: implications for black coral taxonomy and systematics

Although black corals inhabit all the world's oceans, they have been relatively understudied as ～185 of 247 species occur at depths > 50 m. Antipatharians have been included in several phylogenetic studies; however, sample sizes are small and taxonomic coverage minimal. Low levels of mitochondrial (mt) sequence divergence within Scleractinia and Octocorallia are assumed to apply to all anthozoans, although no formal study has been conducted on the order Antipatharia. To quantify genetic variation in the black coral mitogenome, we analysed DNA sequences of the two longest intergenic regions (IGRs) and cox3‐cox1 for 26 of 41 genera, representing all families and subfamilies. We also quantified divergence at the intraspecific level using six mtIGRs and their flanking protein‐coding genes and rRNA for 100+ colonies of Antipathes griggi. Utilizing sequence data from the two mtIGRs, cox3‐cox1, as well as nuclear 18S and 28S, we constructed the first multi‐locus phylogenies of the Antipatharia. Reconstructions revealed that species in the genus Stichopathes are split across two families, Sibopathes macrospina groups among North Atlantic Parantipathes (suggesting the actinopharynx and mesenteries were secondarily lost), and three families are polyphyletic. These and other results provide novel, independent insights into the evolutionary history of antipatharians and support placement of species into higher‐level groupings based on microscopic skeletal features rather than gross colony morphology. An illustrated key to the seven currently recognized families is also provided. © 2013 The Linnean Society of London     

Sexual reproduction and larval settlement of the zooxanthellate coral Alveopora japonica Eguchi at high latitudes

Sexual reproduction and larval settlement of Alveopora japonica Eguchi were studied at its northernmost distribution in Tokyo Bay (34°58'03'N, 139°46'05'E), where the annual sea temperature ranges from 13 to 27 °C. Alveopora japonica is a hermaphroditic brooding coral with oocytes and spermaries developing on separate mesenteries of the polyp. The oocytes first appeared in October, maturing in late August to early September of the following year, whereas the spermaries were first observed in May, and matured in approximately 4 months. The oocytes reached ca. 800 µm in diameter. Planulae containing zooxanthellae in their endoderm were released during the daytime in September and October. Well-developed planulae were able to settle and metamorphose within 7 h. The polyps started budding about 3 weeks after settlement, and took 3 years to grow to maturity. The population examined was sexually reproductive, indicating that A. japonica maintains local populations in Tokyo Bay by sexual reproduction.     

Sexual reproduction and diversity: Connection between sexual selection and biological communities via population dynamics

Sexual reproduction is a mysterious phenomenon. Most animals and plants invest in sexual reproduction, even though it is more costly than asexual reproduction. Theoretical studies suggest that occasional or conditional use of sexual reproduction, involving facultative switching between sexual and asexual reproduction, is the optimal reproductive strategy. However, obligate sexual reproduction is common in nature. Recent studies suggest that the evolution of facultative sexual reproduction is prevented by males that coerce females into sexual fertilization; thus, sexual reproduction has the potential to enforce costs on a given species. Here, the effect of sex on biodiversity is explored by evaluating the reproductive costs arising from sex. Sex provides atypical selection pressure that favors traits that increase fertilization success, even at the expense of population growth rates, that is, sexual selection. The strength of sexual selection depends on the density of a given species. Sexual selection often causes strong negative effects on the population growth rates of species that occur at high density. Conversely, a species that reduces its density is released from this negative effect, and so increases its growth rate. Thus, this negative density-dependent effect on population growth that arises from sexual selection could be used to rescue endangered species from extinction, prevent the overgrowth of common species and promote the coexistence of competitive species. Recent publications on sexual reproduction provide several predictions related to the evolution of reproductive strategies, which is an important step toward integrating evolutionary dynamics, demographic dynamics and community dynamics.     

The social feather duster worm Bispira brunnea (Polychaeta: Sabellidae): aggregations,morphology and reproduction

The fan worm Bispira brunnea is one of the most attractive sabellid polychaetes from Caribbean coral reef areas and it is exploited for ornamental purposes. An understanding of the structure of its aggregations, morphology and reproductive biology will provide information required to facilitate artificial propagation of this species. Ten aggregations were collected in October 2013, February and March 2014 in the Majahual reef lagoon, Mexican Caribbean. Whole aggregations were examined under light microscopy and scanning electron microscopy and the histology of oogenesis was determined. Aggregations were composed of 24–56 individuals and included juveniles and adults. The adults (5–20?mm) did not display any noticeable form of sexual dimorphism. In B. brunnea both sexual (hermaphroditism or gonochorism) and asexual reproduction occurred at the same time within the population: 92.71% reproduced sexually and 52% by architomy. The buds produced by architomy were inside the parental tube, at three regenerative stages. The sex ratio was 36.75% males, 33.11% females, 22.84% hermaphrodites and 7.28% unknown (gametes not seen). Gametes were distributed in the last thoracic segments and throughout the abdomen. Oogenesis was extra-ovarian, development followed four discrete stages and the oocytes were small and asynchronous (60.97 μm in diameter). Sperm morphology was adapted to external fertilization in the water column. Sequential (protandrous) hermaphroditism is suggested to occur in B. brunnea. The pyramidellid ectosymbiont mollusc Odostomia (Eulimastoma) caniculatum is recorded here for the first time as being associated with a sabellid worm.     

Developmental constraints on the mode of reproduction in the facultatively parthenogenetic cockroach Nauphoeta cinerea

Considerable work in evolutionary biology has focused on the question of why sex persists. Both advantages to sex and constraints limiting a return to asexual reproduction are hypothesized to maintain sex once it evolves. Developmental constraints would limit asexual reproduction from a sexual species if it were difficult for females to switch from making eggs that do not develop without fertilization to making zygotes that are capable of developing in the absence of fertilization. Nauphoeta cinerea is an ovoviviparous cockroach in which some females are capable of switching from a sexual mode of reproduction to an asexual mode when isolated from males. Yet, while facultative parthenogenesis can occur in individuals, few females make the switch. Thus, this cockroach provides an ideal system for examining the potential role of developmental constraints in maintaining sex. Here we compare the cytogenetics and embryonic development of sexual and parthenogenetic offspring in N. cinerea. We find that deviations from normal ploidy levels are associated with abnormal development. All viable N. cinerea embryos exhibit typically hemimetabolous insect embryogenesis. Although there is no variation among embryos in development within a sexually produced clutch, we see extreme variation in asexually derived clutches. These results suggest that developmental constraints limit the success of asexual reproduction in this facultatively parthenogenetic cockroach. Our data further suggest that the specific constraint occurs in the switch from a meiotic mode of reproduction requiring fertilization to diploid zygotes that develop in the absence of fertilization.     

Sex allocation in a simultaneous hermaphrodite, the blue-banded goby (Lythrypnus dalli): the effects of body size and behavioral gender and the consequences for reproduction

Although species with both male and female sexual functionsare often dichotomized into simultaneous and sequential hermaphrodites,many simultaneous hermaphrodites also exhibit sequential changesin sex allocation. In a field experiment using one such species,the gobiid fish, Lythrypnus dalli, female-biased individualsreallocated to male function in relation to their relative bodysize: consistent with the sizeadvantage hypothesis, large femaleswere more likely to reallocate and large fish had the highestspawningrates. Individuals, despite internal allocation to bothsexual functions, adopted only one behavioral gender. Behavioralmales had higher reproductive rates than behavioral females,and laboratory experiments showed that females preferred tomate with large males. Behavioral males grew more rapidly anddid notdiffer from behavioral females in survivorship. In addition,individuals who adopted male behavior but did not receive eggsin their nests maintained high levels of female tissue, whereasmales that received eggs did not. Laboratory experiments showedthat, unlike most hermaphroditic animals, L. dalli canchangeallocation either from ‘female’ to ‘male’or from ‘male’ to ‘female’. Thus, L.dalli shares haracteristics of both sequential and simultaneoushermaphrodites. Simultaneous hermaphroditism maybe maintained,in this species, to facilitate rapid sex change from femaleto male and to retain flexibility o that unsuccessful malescan revert to reproduction as females.     

Sexual reproduction,development and larval biology in scleractinian corals

This paper brings together widely scattered information on sexual reproduction in scleractinian corals. It includes a review of information and ideas on sex determination, gametogenesis, gametogenic cycles, fertilization and embryonic development, spawning and planula release, larval behavior, settlement and metamorphosis. The review deals with corals from different habitats and organismic assemblages, including tropical reef corals, temperate water corals, solitary and colonial forms. A summary table of coral species and their known reproductive characteristics is presented.     

Barriers to sexual reproduction in Polygonum viviparum: a comparative developmental analysis of P. viviparum and P. bistortoides

Polygonum viviparum is widely distributed in arctic and alpine regions of the northern hemisphere. Fruit set has never been observed in North American populations and has been reported only very rarely in Europe. Although this species is extremely well studied, the impediments to successful fruit production are unknown. We investigated the sexual reproductive process in P. viviparum growing in the southern Colorado Rocky Mountains. For comparison, we also examined this process in the sympatric congener P. bistortoides, in which reproduction is exclusively sexual. Lack of viable fruit production in P. viviparum has no single developmental explanation; defects occur in each of the processes and structures associated with sexual reproduction studied, yet, these processes and structures also appear to function normally in at least some flowers or individuals. Development is abnormal in many ovules of P. viviparum, however, comparison with P. bistortoides shows that these abnormalities do not contribute to differences in seed production between the two species. The virtual absence of sexual reproduction in P. viviparum appears to be due largely to a low rate of fertilization and to embryo/fruit abortion.     

Elimination of plastids during spermatogenesis and fertilization in the plant kingdom

Ultrastructural and genetic investigations involving diverse species of plants have demonstrated that plastids may be transmitted either biparentally or maternally during sexual reproduction. In species in which plastid transmission is maternal, elimination of plastids from the paternal parent may occur in a number of ways: exclusion from the male gamete during spermatogenesis, loss from the motile sperm, exclusion during fertilization, or degradation within the zygote. These diverse ways in which maternal inheritance of plastids is achieved suggest that this inheritance pattern may have evolved independently many times in response to different selective pressures in different phyletic lineages.     

Developmental flexibility and the effect of social environment on fertility and fecundity in parthenogenetic reproduction

One specialized environment that can influence development arises in the context of social interactions, including the environment contributed by a sexual partner during sexual reproduction. It is often difficult, however, to separate out the effect of mating (fertilization) from the effect of social environment. In the study reported here we examine the effect of social environment mediated by a pheromonal signal on the fertility and fecundity of the facultatively parthenogenetic cockroach Nauphoeta cinerea. By examining parthenogenetically reproducing females, we isolate the effects of social environment in the absence of mating or fertilization. Females exposed to male odors are more likely to produce parthenogenetic offspring. Further, increased exposure to the male pheromone increases the number of offspring produced. Variation in timing of reproduction is also dependent on the male. Thus, social environments are a mechanism by which males contribute to the development of their offspring, resulting in variation in development. This study illustrates the potential evolutionary importance of social environments in development, because a requirement for male-contributed environments may be a constraint to evolving asexual reproduction from a sexually reproducing species.     

Early Life History of the ‘Irukandji’ Jellyfish Carukia barnesi

Adult medusae of Carukia barnesi were collected near Double Island, North Queensland Australia. From 73 specimens, 8 males and 15 females spawned under laboratory conditions. These gametes were artificially mixed which resulted in fertilized eggs. Post fertilization, most eggs developed to an encapsulated planula stage and then paused for between six days and six months prior to hatching as ciliated planulae. The paused stage planulae were negatively buoyant and adhered to substrate. The first planula was produced six days post fertilization, lacked larval ocelli, remained stationary, or moved very slowly for two days prior to metamorphosis into primary polyps. Mature polyps reproduced through asexual reproduction via lateral budding producing ciliated swimming polyps, which in turn settled and developed into secondary polyps. Medusae production for this species was in the form of monodisc strobilation, which left behind polyps able to continue asexual reproduction.     

Planulation,larval biology,and early growth of the deep‐sea soft corals Gersemia fruticosa and Duva florida (Octocorallia: Alcyonacea)

Abstract. Although the reproductive biology and early life‐history stages of deep‐sea corals are poorly understood, such data are crucial for their conservation and management. Here, we describe the timing of larval release, planula behavior, metamorphosis, settlement, and early juvenile growth of two species of deep‐sea soft corals from the northwest Atlantic. Live colonies of Gersemia fruticosa maintained under flow‐through laboratory conditions released 79 planulae (1.5–2.5 mm long) between April and early June 2007. Peak planulation in G. fruticosa coincided with peaks in the chlorophyll concentration and deposition rates of planktic matter. Metamorphosis and settlement occurred 3–70 d post‐release. The eight primary mesenteries typically appeared within 24 h, and primary polyps grew to a height of ～6–10 mm and a stalk diameter of ～1 mm within 2–3 months. Planulae of Duva florida (1.5–2.5 mm long) were extracted surgically from several colonies and were successfully reared in culture. Primary polyps reached a height of ～3–4 mm within 2–3 months. No budding of primary polyps was observed in either species over 11–13 months of monitoring, suggesting a very slow growth rate.     

Double fertilization in Gnetum gnemon (Gnetaceae): its bearing on the evolution of sexual reproduction within the Gnetales and the anthophyte clade

The fertilization process in Gnetum is critical to our understanding of the evolution of sexual reproduction within the Gnetales, a monophyletic group of nonfiowering seed plants that are the closest living relatives to flowering plants. Although much is known about the fertilization process in Ephedra, which is basal within the Gnetales, little is known about sexual reproduction in the derived sister groups Gnetum and Welwitschia. Ovules of Gnetum gnemon were collected at various stages after hand pollination and processed for light, fluorescence, and electron microscopy. Approximately 5 d after pollination, pollen tubes reach sexually mature female gametophytes, which are coenocytic. At that time, a binucleate sperm cell is found within each pollen tube. Within 7 d of pollination, double fertilization events occur when each of two sperm nuclei released from a pollen tube fuses with a separate, undifferentiated female nucleus within the free nuclear female gametophyte, which lacks differentiated egg cells. The products of double fertilization are two viable zygotes; endosperm is not formed. The lack of differentiated egg cells in Gnetum gnemon is unparalleled among land plants and the documentation of a regularly occurring process of double fertilization is congruent with the hypothesis that a rudimentary process of double fertilization evolved in a common ancestor of angiosperms and Gnetales.     

A simple test: evaluating explanations for the relative simplicity of the Edwardsiidae (Cnidaria: Anthozoa)

Many members of the cnidarian subclass Zoantharia (sea anemones, corals, and their allies) pass through a larval stage with eight complete mesenteries and without posterior musculature. This larva is usually transient, developing into an adult with 12 or more mesenteries. The adults of one family of sea anemones, the Edwardsiidae, bear the larval number and arrangement of mesenteries and lack the pedal disc seen in other sea anemones. The morphology of the Edwardsiidae has been interpreted in a number of ways: (1) the Edwardsiidae are the most basal extant zoantharian, having diverged before the evolution of additional mesenteries and basal musculature; (2) they are relatively advanced sea anemones that have secondarily simplified because they burrow in sand or mud rather than attaching to a hard substrate; or (3) edwardsiids are derived anemones that have retained a juvenile morphology through paedomorphosis. Phylogenetic analyses of small subunit ribosomal gene sequences reveal that the Edwardsiidae are derived zoantharians, nested within sea anemones. None of the proposed explanations fully explain the edwardsiid's body plan; edwardsiid anatomy is a mosaic of retained primitive and derived features. The results of the present study provide insight into zoantharian phylogeny and illustrate how phylogenetic tests can be used to study the evolution of cnidarian body plans.     

Hermaphroditism in fishes: an annotated list of species,phylogeny, and mating system

Fewer than 1% of vertebrate species are hermaphroditic, and essentially all of these are fishes. Four types of hermaphroditism are known in fishes: simultaneous (or synchronous) hermaphroditism (SH), protandry (male-to-female sex change; PA), protogyny (female-to-male sex change; PG), and bidirectional sex change (BS or reversed sex change in protogynous species). Here we present an annotated list of hermaphroditic fish species from a comprehensive review and careful re-examination of all primary literature. We confirmed functional hermaphroditism in more than 450 species in 41 families of 17 teleost orders. PG is the most abundant type (305 species of 20 families), and the others are much less abundant, BS in 66 species of seven families, SH in 55 species of 13 families, and PA in 54 species of 14 families. The recently proposed phylogenetic tree indicated that SH and PA have evolved several times in not-closely related lineages of Teleostei but that PG (and BS) has evolved only in four lineages of Percomorpha. Examination of the relation between hermaphroditism type and mating system in each species mostly supported the size-advantage model that predicts the evolution of sequential hermaphroditism. Finally, intraspecific variations in sexual pattern are discussed in relation to population density, which may cause variation in mating system.

     

盐度对沙蜇有性繁殖阶段早期发育的影响

近几十年来,沙蜇的频繁暴发给东亚海域的海洋生态系统带来了广泛影响。在秋季,沙蜇成熟的雌雄水母体在沿岸水域聚集产卵,有性繁殖产生的受精卵发育成新的底栖螅状体,为螅状体种群数量进行补充。河口浅滩海域为沙蜇的繁育地,沿岸盐度较低,在秋季降雨期盐度多变,较低、多变的盐度可能对沙蜇有性繁殖阶段的早期发育产生重要作用,从而影响螅状体种群数量的补充。实验设置了4种不同盐度(15、20、25、30)试验组,在不同盐度下对沙蜇受精卵进行培养,探讨盐度对沙蜇早期发育过程中受精卵、浮浪幼虫发育以及早期螅状体生长及存活的影响。试验结果:沙蜇受精卵胚胎发育的适宜盐度为20,发育基本与盐度25、30同步,盐度15受精卵细胞发育迟缓,发育率显著降低;浮浪幼虫发育适宜盐度为20和25,两组浮浪幼虫附着变态率高于盐度15、30,盐度15时浮浪幼虫活力明显降低、发育迟缓,浮浪幼虫在盐度15时水中存活时间较长可达8 d,但附着时间集中在培养后的3、4天,与其他组相同;早期螅状体幼体适宜盐度为20、25、30,早期螅状体存活率、相对增长率及特定生长率均显著高于盐度15,三组间差异不显著。结果表明,盐度显著影响沙蜇有性繁殖阶段...     

Cycle of gonadal development in Eunicella singularis (Cnidaria: Octocorallia): trends in sexual reproduction in gorgonians

Abstract. Eunicella singularis is a gorgonian species whose members are abundant in hard-bottom sublittoral communities of the Mediterranean Sea. The reproductive biology in this species has been examined to better understand the ability in this species to recover from recent mass mortality events. Eunicella singularis is a stable, gonochoric, iteroparous species that reproduces annually and exhibits a seasonal pattern of gametogenesis characterized by a single annual maturation of the gametes. The sex ratio did not significantly differ from 1:1. Oogenesis lasted 13–17 months, beginning between February and June, and ending with the release of 0.7 planula larvae per polyp between late May and July of the following year. The diameter of mature oocytes ranged 450–860 μm. Spermatogenesis was much shorter than oogenesis and occurred over 5–6 months. Gonadal production of both sexes increased in spring and culminated with the spawning of male colonies in late May–June. Fertilization of oocytes and development of the planula larvae occurred within the polyps of female colonies. Planula release was observed in June and July. The patterns emerging from this and previous studies on sexual reproduction of Mediterranean gorgonians suggest that investment in gonad development appears to be related to resource availability.     

Post-burn reproduction of woody plants in a neotropical savanna: the relative importance of sexual and vegetative reproduction

1. Burning typically occurs at intervals of 1–3-years in the Brazilian cerrado, a rate that exceeds the precolonization fire regime. To determine if woody plants of the cerrado successfully reproduce within the short span of time between burns, experimental burns were used to quantify the effects of fire on sexual and vegetative reproduction of six species of resprouting trees and shrubs.
2. Four of the six species reproduce vegetatively by producing root suckers. For three of these species, Rourea induta , Myrsine guianensis and Roupala montana , sucker production was seven to 15 times greater in burned plots than in unburned controls.
3. Fire had a negative impact on sexual reproduction. Fire caused an immediate reduction in sexual reproductive success by destroying developing reproductive structures and seeds. Additionally, five of the six study species exhibited overall reductions in seed production in the years following fire. Fire had this effect by reducing the individual size of all species and, for three species, by reducing size-specific reproductive output. Only the tree Piptocarpha rotundifolia exhibited increased seed production following burning.
4. Fire caused substantial mortality to both seedlings and suckers. Suckers were larger than seedlings and experienced lower mortality rates for two of three species. Fire-induced mortality of seedlings varied greatly among species, ranging from 33% to 100%.
5. The results indicate that vegetative reproduction is much more successful than sexual reproduction under the high fire frequency typical of current fire regimes. It is concluded that current fire regimes must be causing a shift in species composition, favouring species capable of vegetative reproduction.     

Sexual systems in scleractinian corals: an unusual pattern in the reef-building species Diploastrea heliopora

The sexual system in corals refers to the spatial and temporal pattern of sexual function within an individual coral polyp, colony or population. Although information on sexual systems now exists for over 400 scleractinian species, data are still lacking for some important reef-building taxa. The vast majority of scleractinians are either simultaneous hermaphrodites or gonochoric with other sexual systems rarely occurring. Diploastrea heliopora is one of the most ubiquitous and easily recognised reef-building species in the Indo-West Pacific; however, surprisingly little is known about its reproductive biology. The aim of the present study was to examine the reproductive biology of D. heliopora colonies on chronically impacted, equatorial reefs south of Singapore. Here we show that in Singapore, D. heliopora is a broadcast spawner with predominantly gonochoric polyps. Colonies, however, contained male, female and a low proportion of cosexual polyps during the 14-month sampling period. The most plausible explanation for this is that polyps switch sexes with oogenic and spermatogenic cycles occasionally overlapping. This leads to colony level alternation of sex function within and between breeding seasons. While this sexual system is atypical for scleractinians, it supports molecular evidence that D. heliopora is phylogenetically distinct from species formerly in the family Faviidae.     

Sexual reproduction in the invasive species Fallopia japonica (Polygonaceae)

Fallopia japonica (Polygonaceae) is an invasive perennial plant, well known in North America for its ability to spread aggressively via vegetative reproduction. The contribution of sexual reproduction to the distribution of this species is not well documented, and as a result, F. japonica is treated solely as a clonal species. To investigate the role of sexual reproduction in this species, germination experiments were conducted using seed collected from 29 parents from field sites in Massachusetts and from four greenhouse-grown cultivars. Results showed that wild F. japonica produce large quantities of seed that typically have high germinability. This seed is viable whether sown immediately after collection or subjected to various conditions during the winter season and germinated the following spring. Cultivars of F. japonica also produce viable seed and can thus contribute to the invasiveness of this species. In addition, wild F. japonica seedlings were observed at several field sites, with several of these seedlings surviving the winter and resprouting the following spring. That sexual reproduction and seedling survival occur in the wild has strong implications for the development of management strategies for this species.