Gonochorism vs. hermaphroditism: relationship between life history and fitness in three species of Ophryotrocha (Polychaeta: Dorvilleidae) with different forms of sexuality

1. The relationships between life history, fitness and sexuality, together with their ecological and evolutionary significance, has been analysed comparing the main life-history traits and demography in three closely related species belonging to the genus Ophryotrocha. The species are: the gonochoristic O. labronica, the simultaneous hermaphrodite O. diadema and the protandrous hermaphrodite O. puerilis. 2. Survivorship and reproductive data were collected weekly and were used to construct life tables and population projection matrices for each species and compare life-history characteristics. Elasticity, life-table response and decomposition analyses were performed to examine the relative contribution of fecundity and survivorship to differences in lamda between species. 3. The gonochoristic and hermaphroditic species differ in all the main life-history parameters and also in demographic characteristics. In particular the value of lamda, used commonly to express fitness, is markedly higher in the gonochoristic species while in terms of fitness simultaneous and sequential hermaphroditism are very similar. In the genus Ophryotrocha gonochorism currently represents the most widespread condition, being characteristic of the majority of the known species in the genus. 4. Given the demographic advantage ensured by gonochorism, it remains be understood why some species have retained simultaneous hermaphroditism and one has evolved a sequential type hermaphroditism; the most probable hypothesis is correlated with the density of the species in natural habitats.     

Evaluation of the pair-culture effect inOphryotrocha puerilis (Polychaeta: Dorvilleidae). I. Pair-culture effect and sex ratio

Pairs and larger groups of femaleOphryotrocha puerilis puerilis were formed from formerly isolated specimens. Neither the diameter of the oocytes present in the coelomic fluid nor the number of setigerous segments (ss) of the partners of a newly formed pair allow us to predict which one of the two animals will exhibit sex reversal. Amputation of the palps showed that these ventrolateral appendages of the prostomium are not responsible for the transmission of the mutual influence which is exerted during the pair-culture effect. Isolated females do not produce egg masses but keep their oocytes in the body cavity until they are eventually resorbed. The shedding of oocytes in one of the females of a newly formed pair was formerly considered to be the first step in the pair-culture effect. The present results demonstrate that egg laying in these cases is unspecific and due rather to the end of isolation than to specific interactions with the partner. In groups consisting of up to 50 animals the sex ratio oscillates around 1:1. The presence of primary males does not influence the sex ratio of adult specimens, although these males are capable of fertilizing egg masses produced by adult females. As a result, a high percentage of both young and old males are found in densely populated bowls. The significance of the pair-culture effect in natural populations is discussed in the light of these findings.     

Cytophysiological study of neurosecretory and pheromonal influences on sexual development in Ophryotrocha puerilis (Polychaeta,Dorvilleidae)

Summary

Prominent secretory nerve endings are found at the posterior margin of the supraesophageal ganglion in the protandric polychaete, Ophryotrocha puerilis. Solitary juveniles developing as primary males, and then as females, accumulate neurosecretory material in the nerve endings which thereby swell and become filled with granules. Females maintained in mass culture have similar terminals, whereas in secondary males (males which had been females before), these axon terminals are very small and contain no material. When such males are isolated, they accumulate neurosecretory material within the nerve endings and become females. When formerly isolated females are put together, their stores of neurosecretory material are rapidly discharged. Subsequently they lay egg masses and switch to the male state. These effects are mediated by a pheromone released during social contact of formerly isolated females. The complexity of the relationship between neurosecretory activity and sexual state is indicated by the situation in animals maintained in pairs, when both male and female partners have swollen nerve endings packed with secretory material.     

NOR regions of polychaete worms of the genus Ophryotrocha studied by chromosome banding techniques and FISH

This article reports the results of cytogenetic analyses carried out on 10 species of polychaete worms belonging to the genus Ophryotrocha (Dorvilleidae). Nucleolar organizer regions (NORs) were characterized by Ag staining, C-banding, CMA3 staining, and ribosomal fluorescent in situ hybridization (rDNA FISH). Extensive intraspecific variation in NOR number and distribution were observed in O. costlowi, O. sp. macrovifera, O. notoglandulata, O.l. labronica, O. l. pacifica (2n = 6), O. p. puerilis, O. diadema (2n = 8), O. hartmanni, O. gracilis (2n = 10). In O. sp. robusta (2n = 10), Ag-NORs were always located on a single chromosome pair. CMA3 staining suggests a possible trend toward a GC-rich rDNA compartmentalization. In O.l. labronica, O. p. puerilis, O. diadema, and O. sp. robusta rDNA FISH shows that Ag and FISH signals coincide. Results from C-banding seem to indicate that the increased genome size (GS) observed in O. sp. macrovifera (0.8 pg) and O. hartmanni (1.16 pg) compared to the base GS value of the genus (0.4 pg) cannot be attributed to variation in the heterochromatin content.     

Characterization of novel F-actin envelopes surrounding nuclei during cleavage of a polychaete worm

F-actin accumulations and their possible functions were investigated during cleavage of the polychaete Ophryotrocha puerilis. Unusual cytoplasmic accumulations of F-actin were detected which have never been described before in animal embryos. As shown by TRITC-phalloidin labeling, envelopes of F-actin surrounded late prophase nuclei for a short period of time. DTAF-immunofluorescence of beta-tubulin showed that the F-actin envelope was closely associated with microtubules of the developing spindle apparatus. However, experimental disassembly of microtubules by nocodazole did not prevent the assembly of the F-actin envelope. Disturbance of F-actin envelope formation by cytochalasin B did not alter the course of mitotic events, i.e. position of the nuclei and orientation of the spindle apparatus were not affected, although the respective blastomeres remained uncleaved. However, disassembly of the F-actin envelope correlated temporally with breakdown of the nuclear envelope. Therefore, it is suggested that this new structure plays a role in fragmentation of the nuclear envelope during cleavage of Ophryotrocha puerilis.     

Caddisfly egg mass morphology mediates egg predation: potential costs to individuals and populations

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Cryptic species and colonization processes in Ophryotrocha (Annelida,Dorvilleidae) inhabiting vertebrate remains in the shallow‐water Mediterranean

Over the past several years, there has been growing interest in how bones of decaying mammals are colonized in the marine seabed. One of the most common opportunistic taxa occurring worldwide on bones is dorvilleid polychaetes of the genus Ophryotrocha. In a recent study in the Mediterranean, Ophryotrocha puerilis and Ophryotrocha alborana were two of the most abundant species occurring in experimentally deployed bones. These species have direct development and this makes them a suitable model to study the mechanisms and processes allowing organisms lacking a dispersive larval phase to colonize new substrates. Here, we address the colonization processes at the molecular level for populations of O. puerilis and O. alborana on experimentally deployed mammal bones in the shallow‐water Mediterranean collected over a year at 3‐month intervals. High genetic distances between some of the O. puerilis organisms collected indicated the occurrence of at least two cryptic sibling species (O. puerilis ‘Shallow’ and O. puerilis ‘Deep’) apart from O. puerilis sensu stricto. This was corroborated with phylogenetic analyses using an alignment of three concatenated genes (COI, 16S, H3) and with species delimitation analyses using COI. The haplotype network inferred from COI sequences for O. puerilis ‘Shallow’ showed a few common haplotypes shared between the two trimesters analysed and several other less represented haplotypes only present in one trimester. Thus, colonization of these experimental bones may have been achieved by a few organisms that arrived to the bones and were able to reseed, and by several individuals arriving to the experimental bones and not persisting across time. In contrast, the haplotype network for O. alborana revealed that none of the haplotypes present in three different trimesters were shared, suggesting that the populations arriving at the bones during each trimester were totally replaced by new individuals during the subsequent trimesters. Our study suggests that different species of shallow‐water Ophryotrocha occurring in the Mediterranean may have different patterns of substrate colonization despite sharing similar life histories.     

Molecular phylogeny of the model annelid Ophryotrocha.

Annelids of the genus Ophryotrocha are small opportunistic worms commonly found in polluted and nutrient-rich habitats such as harbors. Within this small group of about 40 described taxa a large variety of reproductive strategies are found, ranging from gonochoristic broadcast spawners to sequential hermaphroditic brooders. Many of the species have a short generation time and are easily maintained as laboratory cultures. Thus they have become a popular system for exploring a variety of biological questions including developmental genetics, ethology, and sexual selection. Despite considerable behavioral, reproductive, and karyological studies, a phylogenetic framework is lacking because most taxa are morphologically similar. In this study we use 16S mitochondrial gene sequence data to infer the phylogeny of Ophryotrocha strains commonly used in the laboratory. The resulting mtDNA topologies are generally well resolved and support a genetic split between hermaphroditic and gonochoristic species. Although the ancestral state could not be unambiguously identified, a change in reproductive strategy (i.e., hermaphroditism and gonochorism) occurred once within Ophryotrocha. Additionally, we show that sequential hermaphroditism evolved from a simultaneous hermaphroditic ancestor, and that characters previously used in phylogenetic reconstruction (i.e., jaw morphology and shape of egg mass) are homoplasic within the group.     

Egg trading in the simultaneously hermaphroditic polychaete worm Ophryotrocha gracilis (Huth)

Ophryotrocha gracilis (Huth 1934) is a simultaneously hermaphroditicpolychaete worm with external fertilization and a brief protandrousphase. The mating system of this species seems to meet conditionsleading to the establishment of egg-trading behavior. Experimentsshowed that mating occurs in pairs composed of two simultaneoushermaphrodites; sex roles are sequentially alternated and self-fertilizationis avoided. Egg reciprocation is kept evolutionarily stableby laying eggs in multiple, small-sized egg clutches and reducingthe reproductive success of pairs in which one of the two partnersdoes not reciprocate egg exchange. The frequency of ovigeroushermaphrodites in mass cultures of O. gracilis is about 50".Such a high mate availability preadapts hermaphrodites of O.gracilis to change partners very frequently and to reduce investmentin parental care, contrary to what is observed in another egg-trading,simultaneously hermaphroditic species, O. diadema. Laboratorypopulations of O. diadema have a frequency of only 17% ovigeroushermaphrodites     

Systematics and biodiversity of Ophryotrocha (Annelida,Dorvilleidae) with descriptions of six new species from deep-sea whale-fall and wood-fall habitats in the north-east Pacific

Six new Ophryotrocha species are described from five whale-falls and two wood-falls off the southern Californian coast. Phylogenetic analyses based on the nuclear gene H3 and the mitochondrial genes COI and 16S using MrBayes and maximum likelihood analyses were performed on 40 dorvilleid taxa and one outgroup. Ophryotrocha batillus sp. nov. is morphologically identical to Ophryotrocha scutellus described from a shallow water whale-fall in the North Atlantic, although the two cryptic species differ genetically. Ophryotrocha langstrumpae sp. nov. is closely related in the molecular phylogenetic analyses to these two sibling species. Ophryotrocha flabella sp. nov. is similar to Ophryotrocha globopalpata, and although there are a few morphological differences, the genetic divergence is low between the two species. Ophryotrocha nauarchus sp. nov. is sexually dimorphic, with males having appendages on the first chaetiger. Ophryotrocha magnadentata sp. nov. and Ophryotrocha longicollaris sp. nov. are sister species in our molecular analyses, and together with O. nauarchus sp. nov. and O. flabella sp. nov. they fall within a clade that includes O. globopalpata and Exallopus jumarsi described from hydrothermal vents in the Pacific Ocean and Ophryotrocha longidentata from the shallow North Atlantic. Our results highlight the remarkable unknown diversity of deep-water habitats and the role of chemosynthetic ecosystems in the evolution of deep-sea life.     

Jaw morphology and ontogeny in five species of Ophryotrocha

Detailed scanning electron microscopy of jaws within the genus Ophryotrocha (Dorvilleidae, Annelida) was performed on 871 jaw parts. The investigations resulted in new understandings of the ontogeny and jaw morphology and have systematic implications for the family. Five species in the genus (Ophryotrocha alborana, O. diadema, O. gracilis, O. hartmanni, and O. labronica pacifica) were kept in culture, and the development of the jaws was studied by sampling throughout their life history. Ophryotrocha species have mandibular plates that remain the same throughout ontogeny, whereas the posterior shafts elongate. Both mandibular plate morphology and shaft ontogeny have species‐specific distinctions. In Ophryotrocha, the maxillae can be assigned to three to four distinct types, which are replaced by moulting. The maxillary morphology and developmental stages at which moults occur are species specific, although with broad intervals. A redefinition is given for some of the basic jaw elements, and new homologies are proposed for structures that are also present across other dorvilleid taxa. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Genome size and karyotype length in some interstitial polychaete species of the genus Ophryotrocha (Dorvilleidae).

Interstitial polychaetes of the genus Ophryotrocha are very small, progenetic, and morphologically very similar. These worms have been widely used in evolutionary biology and sexuality studies. To have a better insight into the karyological evolution of this genus, we measured the total karyotypic length and the 2C nuclear DNA content of the nine best-known species of this genus. No interspecific differences were observed in karyotypic lengths, apart from that of O. gracilis, which was significantly greater than the karyotypic length of five of the nine species. The genome size (i.e., 1C DNA content calculated from 2C DNA content) in eight of the nine species is about 0.4 pg, irrespective of the chromosome number. A group of four gonochoric and morphologically indistinguishable species, with 2n = 6 metacentric chromosomes, appears to be heterogeneous with regard to its DNA content, because one of the species, O. macrovifera, has a genome twice the size of that of the other three species. A hermaphroditic species, O. hartmanni, has a genome three times that size. No correlation has been observed between genome size and body size, egg cell diameter, or time interval from egg fertilization to sexual maturity. The basic genome size of 0.4 pg is among the lowest recorded in invertebrates. Hypotheses about selective pressures that maintain such a low amount of nuclear DNA in this genus are discussed.     

Late Palaeozoic mollusc reproduction: cephalopod egg-laying behavior and gastropod larval palaeobiology

Faunal analysis of an oxygen‐depleted marine Lower Carboniferous succession (Late Mississippian Ruddle Shale) suggests how some cephalopod taxa laid their eggs during the Late Palaeozoic. At the Ruddle Shale collecting site in Arkansas, USA, the facies and overall fauna suggest severe oxygen depletion at the sediment/water interface. The Ammonoidea, with their small egg size, were probably laid in suspended gelatinous egg‐filled masses in the water column above the bottom or by attachment of the egg masses to floating debris. The ammonitella embryos developed within the suspended or attached egg mass; hatched individuals became part of the free‐swimming plankton biota. Based on shell morphology the Bactritoidea probably followed the same reproductive pattern. Coiled nautiloids (the Nautilida) and most orthoconic nautiloids (mostly the Pseudorthocerida) probably did not lay their eggs in the mid water column or as floatant attachments. This conclusion is based on the fact that, with one exception, all shells recovered of these two nautiloid orders are well past hatching. Gastropods in the Ruddle Shale are very small and cannot be visually detected in the field. However, microgastropods are abundant in washed residues. Most specimens are much smaller than 1 mm. The largest caenogastropod specimen is 1.3 mm high. These caenogastropods represent isolated larval shells and a successful metamorphosis was impossible because of oxygen depletion on the bottom. Allegations that a size of more than 1 mm is too large for pelagic larvae are refuted by examples of planktotrophic larval shells of modern gastropods (more than 1 mm high) and Triassic caenogastropods (up to 2 mm high) from the Cassian Formation (Northern Italy, South Alps). Repository information is given for the type‐material of the gastropod species Nuetzelina striata  Bandel, 2002 and Anozyga arkansasensis  Bandel, 2002 which were both erected based on specimens from the Ruddle Shale that were illustrated by Nützel & Mapes in 2001.     

The survival of Ostertagia circumcincta and Trichostrongylus colubriformis in faecal culture as a source of bias in apportioning egg counts to worm species.

When cultured alone or concurrently with Trichostrongylus colubriformis in sheep faeces, Ostertagia circumcincta produced fewer infective larvae per 100 eggs than did T. colubriformis. Averaged over five trials 60% of T. colubriformis eggs were recovered as infective larvae while for O. circumcincta the figure was only 39%. This result was observed for two strains of O. circumcincta and was independent of when larvae were harvested from culture (days 6-10 at 25 degrees C). The mortalities of both species occurred at the first and second larval stages. These observations are of concern when using larval differentiation from faecal culture to make quantitative estimates of worm egg numbers for each species present. Species such as T. colubriformis which have a low mortality during culture are likely to have their egg numbers overestimated when cultured with a species, like O. circumcincta, that suffers high mortality in culture.     

A REVIEW OF LARVAL DEVELOPMENT IN THE INTERTIDAL LIMPET GENUS SIPHONARIA (GASTROPODA: PULMONATA)

A compilation of distributional and life-history data relatingto mode of larval development is presented for 26 species ofSiphonana, a genus of intertidal pulmonates. Most species depositgelatinous benthic egg masses with two species releasing pelagicegg masses. Fifteen species hatch as planktonic-developing larvae,nine hatch as direct-developing juveniles, and in a furthertwo larvae hatch with both the swimming velar apparatus (associatedwith plank-tonic development) and a crawling foot (associatedwith direct development). Data on mode of larval developmentare interpreted with respect to some adaptive models. Despiteimportant exceptions, there is support for adaptive models basedupon egg capsule size (direct developers hatch from larger eggcapsules) and intertidal distribution (direct developers generallyoccur higher on the shore than planktonic developers). Worldwide,planktonic developers are more widespread than direct-developingspecies, and individual planktonic species have a greater meanlatitudinal range. The evidence for adaptive models relatinglatitudinal distribution to developmental mode is equivocal.There appears to be no clear relationship between body sizeand developmental mode in the genus, although the smallest specieshas direct development and the largest has planktonic development.In most siphonariid subgenera, developmental mode appears tobe constant, but two subgenera contain a mixture of developmentaltypes (Received 1 November 1993; accepted 15 April 1994)     

DEVELOPMENTAL CONSEQUENCES OF AN EVOLUTIONARY CHANGE IN EGG SIZE: AN EXPERIMENTAL TEST

Larvae of two species of sea urchins (Strongylocentrotus droebachiensis and S. purpuratus) differ in initial form and in the rate of development. To determine whether these differences are attributable to the large interspecific difference in egg size, we experimentally reduced egg size by isolating blastomeres from embryos. The rate of development of feeding larvae derived from isolated blastomeres was quantified using a novel morphometric method. If the differences early in the life histories of these two species are due strictly to differences in egg size, then experimental reduction of the size of S. droebachiensis eggs should yield an initial larval form and rate of development similar to that of S. purpuratus. Our experimental manipulations of egg size produced three clear results: 1) smaller eggs yielded larvae that were smaller and had simpler body forms, 2) smaller eggs resulted in slower development through the early feeding larval stages, and 3) effects of egg size were restricted to early larval stages. Larvae from experimentally reduced eggs of the larger species had rates of development similar to those of the smaller species. Thus, cytoplasmic volumes of the eggs, not genetic differences expressed during development, account for differences in larval form and the rate of form change. This is the first definitive demonstration of the causal relationship between egg size (parental investment per offspring) and life-history characteristics in marine benthic invertebrates. Because larval form influences feeding capability, the epigenetic effects of egg size on larval form are likely to have important functional consequences. Adaptive evolution of egg size may be constrained by the developmental relationships between egg size and larval form: evolutionary changes in egg size alone can result in concerted changes in larval form and function; likewise evolutionary changes in larval form and function can be achieved through changes in egg size. These findings may have broader implications for other taxa in which larval morphology and, consequently, performance may be influenced by changes in egg size.     

Effects of temperature on hatching time and hatchling proportions in a poecilogonous population of Haminoea zelandiae

Poecilogony is a relatively uncommon life-history strategy that results in the production of two different larval forms from the same egg mass (e.g., free-swimming lecithotrophic larvae and post-metamorphic, crawling juveniles). In this study, a population of the opisthobranch gastropod Haminoea zelandiae from Pauatahanui Inlet, New Zealand, was found to exhibit poecilogony. Further, differences in development, hatching times and proportion of hatchlings that were veligers or juveniles were examined for egg masses in two temperature regimes in the laboratory: cool (15-17 °C), and warm (21-23 °C). Hatching proportions were also examined for egg masses collected from the field (where temperatures ranged from 21-23 °C) for varying lengths of time (1 d, 5 d, and 10 d post-spawning). Hatchlings from egg masses in warmer temperatures developed faster and hatched earlier than those in cool temperatures. In the laboratory, egg masses in warm conditions hatched a greater proportion of post-metamorphic juveniles (45.4%) compared to egg masses in cool conditions (24.6%) Further, egg masses that had been in the field 10 d before hatching (i.e., more days at warmer temperatures) exhibited a greater proportion of post-metamorphic juveniles (67.9%) than those that were collected after only 1 d in the field (25.1%). Together these results suggest that temperature may have an important role in mediating dispersal strategies in this poecilogonous species.     

The protective role of endogenous bacterial communities in chironomid egg masses and larvae

Insects of the family Chironomidae, also known as chironomids, are distributed worldwide in a variety of water habitats. These insects display a wide range of tolerance toward metals and organic pollutions. Bacterial species known for their ability to degrade toxicants were identified from chironomid egg masses, leading to the hypothesis that bacteria may contribute to the survival of chironomids in polluted environments. To gain a better understanding of the bacterial communities that inhabit chironomids, the endogenous bacteria of egg masses and larvae were studied by 454-pyrosequencing. The microbial community of the egg masses was distinct from that of the larval stage, most likely due to the presence of one dominant bacterial Firmicutes taxon, which consisted of 28% of the total sequence reads from the larvae. This taxon may be an insect symbiont. The bacterial communities of both the egg masses and the larvae were found to include operational taxonomic units, which were closely related to species known as toxicant degraders. Furthermore, various bacterial species with the ability to detoxify metals were isolated from egg masses and larvae. Koch-like postulates were applied to demonstrate that chironomid endogenous bacterial species protect the insect from toxic heavy metals. We conclude that chironomids, which are considered pollution tolerant, are inhabited by stable endogenous bacterial communities that have a role in protecting their hosts from toxicants. This phenomenon, in which bacteria enable the continued existence of their host in hostile environments, may not be restricted only to chironomids.     

Oviposition in three species of limnephiloid caddisflies (Trichoptera): hierarchical influences on site selection

1. Oviposition site selection was studied in three trichopteran species; an undescribed species of Hydatophylax (Limnephilidae), Onocosmoecus unicolor (Limnephilidae) and Neophylax rickeri (Uenoidae), in two Coastal Range streams in California, U.S.A. 2. Hydatophylax sp. egg masses were generally found at or above the water surface on substrata near the bank in pools, where undercut banks, overhanging vegetation and rocks shaded attachment sites and provided wind protection. Onocosmoecus unicolor females deposited their egg masses exclusively above the water on moist emergent wood. The egg masses of N. rickeri were almost exclusively found in high velocity areas of riffles, where females oviposited under water and attached their egg masses to the submersed undersides of unembedded, protruding stones with large emergent surfaces. 3. For Hydatophylax sp. and N. rickeri, a hierarchical selection scheme is proposed in which females use cues at three different spatial levels (stream, stream subunit, substratum) to choose oviposition sites. Females of O. unicolor seemed to choose at only two different levels of spatial resolution (stream, substratum). 4. The formation of large aggregations of egg masses in N. rickeri and O. unicolor suggests that females actively choose oviposition sites where conspecific egg masses are already attached. The clustering of egg masses may minimise the risk of predation through the dilution effect, because egg masses of N. rickeri and O. unicolor are both attacked by dipteran predators. 5. Females of the three Trichoptera species studied are able to make a clear choice of oviposition site in the heterogeneous stream environment. In comparison with other stream microhabitats, these are characterised by stable and relatively predictable environmental conditions during the time of egg development, which is seen as the main selective pressure leading to the observed egg‐laying behaviour.