Major radiation of cheilostome bryozoans: Triggered by the evolution of a new larval type?

A macroevolutionary model is developed to account for the “adaptive radiation”; of cheilostome bryozoans that commenced in the Cenomanian after a long phase of low diversity. Living cheilostome species possess one of two types of larvae; planktotrophic (cyphonautes) larvae of relatively long duration, and brooded non‐planktotrophic (coronate) larvae of short duration. Planktotrophic larvae characterize the paraphyletic “malacostegans”; from which “advanced”; cheilostomes with non‐planktotrophic larvae are thought to have evolved monophyletically. Research on other marine invertebrates suggests that gene flow within and between populations is likely to be poorer in species having non‐planktotrophic larvae, and hence the frequency of allopatric and quasi‐sympatric speciation may be greater. Skeletal evidence of larval brooding in the cheilostomes first appears in the late Albian, immediately before their adaptive radiation, and the evolution of non‐planktotrophy with associated increase in speciation rate is proposed to have triggered this radiation.     

Crossover distribution and high interference for both the X chromosome and an autosome during oogenesis and spermatogenesis in Caenorhabditis elegans

Regulation of both the number and the location of crossovers during meiosis is important for normal chromosome segregation. We used sequence-tagged site polymorphisms to examine the distribution of all crossovers on the X chromosome during oogenesis and on one autosome during both oogenesis and spermatogenesis in Caenorhabditis elegans. The X chromosome has essentially one crossover during oogenesis, with only three possible double crossover exceptions among 220 recombinant X chromosomes. All three had one of the two crossovers in the same chromosomal interval, suggesting that crossovers in that interval do not cause interference. No other interval was associated with double crossovers. Very high interference was also found on an autosome during oogenesis, implying that each chromosome has only one crossover during oogenesis. During spermatogenesis, recombination on this autosome was reduced by approximately 30% compared to oogenesis, but the relative distribution of the residual crossovers was only slightly different. In contrast to previous results with other autosomes, no double crossover chromosomes were observed. Despite an increased frequency of nonrecombinant chromosomes, segregation of a nonrecombinant autosome during spermatogenesis appears to occur normally. This indicates that an achiasmate segregation system helps to ensure faithful disjunction of autosomes during spermatogenesis.     

Gametogenesis in placental and non-placental ovicellate cheilostome Bryozoa

Oogenesis is compared in two cheilostome bryozoans with contrasting reproductive strategies. from southern Britain: Chartella papyracea (Ellis & Solander) is a non–placental ovicellate brooder, whereas Bugula flabellata (Thompson in Gray) is a placental brooder. The ovarian cycles are similar, and each oocyte develops in tandem with a single nurse cell. Eggs of both species are telolecithal, However, those of B. flabellata are less than 20% the volume of those of the other species, and there are considerable differences in the ultra-structure of oogenesis. In both cases, spermatogenesis has the typical bryozoan pattern. Precocious insemination of the oocyte occurs in both species.     

Comparative anatomical study of internal brooding in three anascan bryozoans (Cheilostomata) and its taxonomic and evolutionary implications

The anatomical structure of internal sacs for embryonic incubation was studied using SEM and light microscopy in three cheilostome bryozoans-Nematoflustra flagellata (Waters,1904), Gontarella sp., and Biflustra perfragilis MacGillivray, 1881. In all these species the brood sac is located in the distal half of the maternal (egg-producing) autozooid, being a conspicuous invagination of the body wall. It consists of the main chamber and a passage (neck) to the outside that opens independently of the introvert. There are several groups of muscles attached to the thin walls of the brood sac and possibly expanding it during oviposition and larval release. Polypide recycling begins after oviposition in Gontarella sp., and the new polypide bud is formed by the beginning of incubation. Similarly, polypides in brooding zooids degenerate in N. flagellata and, sometimes, in B. perfragilis. In the evolution of brood chambers in the Cheilostomata, such internal sacs for embryonic incubation are considered a final step, being the result of immersion of the brooding cavity into the maternal zooid and reduction of the protecting fold (ooecium). Possible reasons for this transformation are discussed, and the hypothesis of Santagata and Banta (Santagata and Banta1996) that internal brooding evolved prior to incubation in ovicells is rejected.     

The Fog-3 Gene and Regulation of Cell Fate in the Germ Line of Caenorhabditis Elegans

In the nematode Caenorhabditis elegans, germ cells normally adopt one of three fates: mitosis, spermatogenesis or oogenesis. We have identified and characterized the gene fog-3, which is required for germ cells to differentiate as sperm rather than as oocytes. Analysis of double mutants suggests that fog-3 is absolutely required for spermatogenesis and acts at the end of the regulatory hierarchy controlling sex determination for the germ line. By contrast, mutations in fog-3 do not alter the sexual identity of other tissues. We also have characterized the null phenotype of fog-1, another gene required for spermatogenesis; we demonstrate that it too controls the sexual identity of germ cells but not of other tissues. Finally, we have studied the interaction of these two fog genes with gld-1, a gene required for germ cells to undergo oogenesis rather than mitosis. On the basis of these results, we propose that germ-cell fate might be controlled by a set of inhibitory interactions among genes that specify one of three fates: mitosis, spermatogenesis or oogenesis. Such a regulatory network would link the adoption of one germ-cell fate to the suppression of the other two.     

Reproduction and developmental pathways of Red Sea Xeniidae (Octocorallia,Alcyonacea)

This work on Red Sea alcyonaceans describes the reproductive patterns of 21 xeniid species. Gonochorism is the commonest sexual mode but simultaneous hermaphroditism was recorded in 4 species and brooding of planulae was observed in 15 species. The reproductive patterns of Xenia umbellata and Heteroxenia fuscescens were examined. X. umbellata exhibits seasonal spermatogenesis, continuous oogenesis and a 7 month period of planulation each year. H. fuscescens has continuous gametogenesis, and planulation occurs throughout the year, lacking any lunar pattern. The prolonged breeding season of H. fuscescens reflects intrapopulation asynchrony in larval development and subsequent maturation. Xeniids possess structurally similar gonads, but adopt diverse reproductive and developmental pathways.     

Comparative anatomy of internal incubational sacs in cupuladriid bryozoans and the evolution of brooding in free‐living cheilostomes

Numerous gross morphological attributes are shared among unrelated free‐living bryozoans revealing convergent evolution associated with functional demands of living on soft sediments. Here, we show that the reproductive structures across free‐living groups evolved convergently. The most prominent convergent traits are the collective reduction of external brood chambers (ovicells) and the acquisition of internal brooding. Anatomical studies of four species from the cheilostome genera Cupuladria and Discoporella (Cupuladriidae) show that these species incubate their embryos in internal brooding sacs located in the coelom of the maternal nonpolymorphic autozooids. This sac consists of a main chamber and a narrow neck communicating to the vestibulum. The distal wall of the vestibulum possesses a cuticular thickening, which may further isolate the brood cavity. The presence of this character in all four species strongly supports grouping Cupuladria and Discoporella in one taxon. Further evidence suggests that the Cupuladriidae may be nested within the Calloporidae. Based on the structure of brooding organs, two scenarios are proposed to explain the evolution of the internal brooding in cupuladriids. The evolution of brood chambers and their origin in other free‐living cheilostomes is discussed. Unlike the vast majority of Neocheilostomina, almost all free‐living cheilostomes possess nonprominent chambers for embryonic incubation, either endozooidal and immersed ovicells or internal brooding sacs, supporting the idea that internal embryonic incubation is derived. We speculate that prominent skeletal brood chambers are disadvantageous to a free‐living mode of life that demands easy movement through sediment in instable sea‐floor settings. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

THE EVOLUTION OF OVIPARITY WITH EGG GUARDING AND VIVIPARITY IN LIZARDS AND SNAKES: A PHYLOGENETIC ANALYSIS

This paper investigates the evolution of viviparity and of egg guarding in lizards and snakes in which three modes of reproduction can be described: oviparity without egg guarding, oviparity with egg guarding, and viviparity. All possible transitions of reproductive modes were detected in each taxon using Maddison's method. We then tested two specific hypotheses. First, egg guarding can be regarded as an alternative to viviparity. A relatively frequent association of egg guarding and viviparous species in the same taxon may be due to similar environmental conditions or species characteristics leading to two different solutions. Second, egg guarding may facilitate the evolution of viviparity. This hypothesis is supported by the high frequency of viviparous species in taxa containing egg guarding species and by a tendency for prolonged uterine retention of eggs in brooding squamates. Our analyses demonstrate that the first hypothesis is the best supported. Egg guarding and viviparity most often evolved independently. If a major benefit of egg guarding is the repulsion of potential predators, size is one of the most obvious morphological characters that should be correlated with the evolution of reproductive modes. The two reproductive traits were correlated to a reduction in body size for viviparous species and an increase in body size for egg guarding species. This could partly explain why the evolution of these reproductive modes seems almost antagonist.     

Male pregnancy in seahorses and pipefish: beyond the mammalian model

Pregnancy has been traditionally defined as the period during which developing embryos are incubated in the body after egg-sperm union. Despite strong similarities between viviparity in mammals and other vertebrate groups, researchers have historically been reluctant to use the term pregnancy for non-mammals in recognition of the highly developed form of viviparity in eutherians. Syngnathid fishes (seahorses and pipefishes) have a unique reproductive system, where the male incubates developing embryos in a specialized brooding structure in which they are aerated, osmoregulated, protected and likely provisioned during their development. Recent insights into physiological, morphological and genetic changes associated with syngnathid reproduction provide compelling evidence that male incubation in these species is a highly specialized form of reproduction akin to other forms of viviparity. Here, we review these recent advances, highlighting similarities and differences between seahorse and mammalian pregnancy. Understanding the changes associated with the parallel evolution of male pregnancy in the two major syngnathid lineages will help to identify key innovations that facilitated the development of this unique form of reproduction and, through comparison with other forms of live bearing, may allow the identification of a common set of characteristics shared by all viviparous organisms.     

Viviparity in the sea star Cryptasterina hystera (Asterinidae)--conserved and modified features in reproduction and development

Cryptasterina hystera has a highly derived life history with intragonadal development and juveniles that emerge from the parent's reproductive tract. The gonads are ovotestes with developing eggs separated from sperm by follicle cells. C. hystera has typical echinosperm that must enter the gonoduct of conspecifics to achieve fertilization. During oogenesis, an initial period of yolk accumulation is followed by hypertrophic lipid deposition, the major contributor to the increase in egg size. 1-Methyladenine induces egg maturation and ovulation, but the spawning component of the hormonal cascade is suppressed. This is the major alteration in reproduction associated with evolution of viviparity in C. hystera. The switch to viviparity was not accompanied by major change in gonad structure, indicating there were few or no anatomical constraints for evolution of a marsupial function for the gonad. Despite their intragonadal habitat, the brachiolaria are equipped for a planktonic life, swimming in gonadal fluid. During the gastrula stage, lipid provisions are released into the blastocoel where they are stored for juvenile development. The eggs of C. hystera have light and dark cytoplasmic regions that mark animal-vegetal polarity. The dark pigment provided a marker to follow the fate of vegetal cells. Live birth is rare in the Echinodermata and the incidence of this form of brooding in the phylum is reviewed.     

Complex heterochrony underlies the evolution of Caenorhabditis elegans hermaphrodite sex allocation

Hermaphroditic organisms are key models in sex allocation research, yet the developmental processes by which hermaphrodite sex allocation can evolve remain largely unknown. Here we use experimental evolution of hermaphrodite‐male (androdioecious) Caenorhabditis elegans populations to quantify the developmental changes underlying adaptive shifts in hermaphrodite sex allocation. We show that the experimental evolution of increased early‐life self‐fertility occurred through modification of a suite of developmental traits: increased self‐sperm production, accelerated oogenesis and ovulation, and increased embryo retention. The experimental evolution of increased self‐sperm production delayed entry into oogenesis—as expected, given the sequentially coupled production of self‐spermatogenesis and oogenesis. Surprisingly, however, delayed oogenesis onset did not delay reproductive maturity, nor did it trade‐off with gamete or embryo size. Comparing developmental time dynamics of germline and soma indicates that the evolution of increased sperm production did not delay reproductive maturity due to a globally accelerated larval development during the period of self‐spermatogenesis. Overall, heterochrony in gametogenesis and soma can explain adaptive shifts in hermaphrodite sex allocation.     

Reproduction and development ofSebastes in the context of the evolution of piscine viviparity

Synopsis Selected aspects of the reproduction and development ofSebastes and other rockfishes are reviewed in the context of piscine viviparity. Among the eight subfamilies of the Scorpaenidae, viviparity is confined to the subfamily Sebastinae; gestation is lumenal and the embryos usually develop to term within the egg envelope. Transitional states from oviparity to viviparity are evident in different species within the family. A scenario for the evolutionary origin of viviparity in rockfishes is derived from an analysis of scorpaeniform reproductive biology. Although viviparity is best developed in the genusSebastes, it is still in a primitive, unspecialized state. Rockfish viviparity is essentially lecithotrophic, i.e. embryonic nutrition is dependent on the energy reserves laid down during oogenesis. In other groups of viviparous fishes, lecithotrophy has been shown to be better suited energetically to seasonally unpredictable habitats, whereas matrotrophy requires a predictable food supply. During the evolution of an essentially primitive form of lecithotrophic viviparity in rockfishes, the advantages of high fecundity associated with oviparity were retained while an enormous increase in the survival rate of the developing embryos was acquired. The basic lecithotrophic pattern of oviparous development was not changed since it offered selective advantages both in terms of energetics and as a basis for retaining a large brood size.     

Biochemical dynamics of spermatogenesis and oogenesis in Eledone cirrhosa and Eledone moschata (Cephalopoda: Octopoda)

The effect of spermatogenesis and oogenesis on protein, lipid, glycogen, cholesterol and energy contents, total amino acid and fatty acid profiles of Eledone cirrhosa and Eledone moschata tissues (gonad, digestive gland and muscle) was investigated. A significant (p<0.05) increase in the amino acids and protein content of the gonad throughout sexual maturation (namely in oogenesis) was observed, but the allocation of these nitrogen compounds from the digestive gland and muscle was not evident. The major essential amino acids (EAA) in the three tissues were leucine, lysine and arginine. The major nonessential amino acids (NEAA) were glutamic acid, aspartic acid and alanine. A significant increase in lipid and fatty acid contents of gonad and digestive gland was observed. There was also little evidence of accumulated lipid storage reserves being used for egg production. It seems that for egg production Eledone species use energy directly from food, rather than from stored products. Most of saturated fatty acid (SFA) content of the three tissues was presented as 16:0 and 18:0, monounsaturated fatty acid (MUFA) content as 18:1 and 20:1 and polyunsaturated fatty acid (PUFA) content as 20:4n-6, 20:5n-3 and 22:6n-3. Cholesterol and glycogen contents significantly increased in gonad and digestive gland throughout maturation while the muscle revealed no obvious pattern. If Eledone's component sterols are of a dietary origin, a considerable variation in the cholesterol content between species might be expected on the basis of the sterol composition of their prey. Although spermatogenesis and oogenesis had a significant effect (p<0.05) in gonad and digestive gland energy content, the biochemical composition of digestive gland and muscle may not be primarily influenced by sexual maturation, but rather by other biotic factors such as feeding activity, food availability, spawning and brooding.     

Hormonal Control of Gametogenesis in the Terrestrial Snail, Euhadra peliomphala

Effects of the cerebral ganglion (brain) and the optic gland on oogenesis and spermatogenesis were studied in the terrestrial snail, Euhadra peliomphula. Removal of the optic tentacles inhibited both oogenesis and spermatogenesis. These effects were abolished by the injection of the optic tentacle extract. In the tissue of the optic tentacle, only the optic gland showed the recovery action. Furthermore, in vitro the optic gland extract stimulated spermatogenesis but had no effect on oogenesis. On the contrary, the brain extract promoted oogenesis not only in vitro and but also in viva The hermaphroditic gland extract reduced the nuclear volume of the optic gland cells, but the brain extract did not. In addition, among sex steroid hormones secreted from the hermaphroditic gland, only testosterone also reduced the nuclear volume of optic gland cells. On the basis of these results, hormonal mechanisms of gametogenesis in Euhadra peliomphala are discussed.     

MOLECULAR PHYLOGENETIC ANALYSIS OF LIFE-HISTORY EVOLUTION IN ASTERINID STARFISH

We analyzed phylogenetic relationships among 12 nominal species of starfish in the genera Patiriella and Asterina (Order Valvatida, Family Asterinidae), based on complete sequences for a mitochondrial protein coding gene (cytochrome oxidase subunit I) and five mitochondrial transfer RNA genes (alanine, leucine, asparagine, glutamine, and proline) (1923 bp total). The resulting phylogeny was used to test a series of hypotheses about the evolution of life-history traits. (1) A complex, feeding, planktonic larva is probably ancestral for these starfish, but this is not the most parsimonious reconstruction of ancestral larval states. (2) The feeding larval form was lost at least four times among these species, and three of these losses occurred among members of a single clade. (3) Small adult size evolved before both cases of hermaphroditism and viviparous brooding, but viviparity was not always preceded by an intermediate form of external brooding. (4) An ordered transformation series from feeding planktonic development to viviparous brooding has been predicted for starfish, but we could not find an example of this transformation series. (5) Viviparity evolved recently (< 2 Mya). (6) Both species selection and transformation of lineages may have contributed to the accumulation of species with nonfeeding development among these starfish. (7) Neither Asterina nor Patiriella are monophyletic genera. Larval forms and life-history traits of these starfish have evolved freely under no obvious constraints, contrary to the widely assumed evolutionary conservatism of early development.     

Reproductive adaptations of Antarctic benthic invertebrates

The majority of Antarctic benthic invertebrates so far studied do not produce pelagic larvae, but develop non-pelagically by means of egg capsules, brooding or viviparity. The predominance of protected development in the Antarctic benthos is primarily due to the short period of summer phytoplankton abundance and the low sea temperature. Such conditions make it difficult for a larva to complete pelagic development before food becomes scarce in the surface waters. Prosobranch gastropods illustrate some important aspects of Antarctic benthic invertebrate reproduction. Species which develop non-pelagically have an aseasonal or prolonged spawning period. They produce a small number of large yolky eggs which remain in the benthos and develop slowly, giving rise to large, fully competent juveniles. Conversely, one species with free development has a short, synchronous spawning period during early summer, producing larvae which can benefit from the phytoplankton bloom. Protected development by means of brooding will limit dispersion, but transport on floating algae and by anchor ice may partially compensate for this in the Antarctic.     

Progress on reproductive modes of sarcosaphagous flies

The reproduction strategies of invertebrates, oviparity, ovoviviparity and viviparity, always reflect the relationship between individuals and their surroundings. There is plasticity in the reproductive strategies of sarcosaphagous flies as they adjust to rapidly changing circumstances. The transition from oviparity to ovoviviparity or viviparity involves numerous changes in physiology, morphology and immunology. Demonstrating these processes can make the application of entomology work in forensic practice more reliable. This essay reviews means of reproduction in sarcosaphagous flies and identifies related features. It is shown that not only the reproduction traits, such as fast location of carrion, and uncommon number of ovaries and oogenesis, but also some morphological features are related to viviparity. In general, viviparous flies have larger adult bodies but smaller eggs and chorions. Moreover, the length of terminalia and the shape of the vagina also varies among those three modes. Reproductive plasticity is a bridge between the three reproductive modes, and it can greatly influence the inference of the post-mortem interval (PMImin).     

Evolutionary origins of viviparity in Chamaeleonidae

Historically, an understanding of viviparity and its evolution in Old World chameleons (Chamaeleonidae) has lagged behind that of other squamate families. Not only is reproductive information scarce or entirely absent for most chameleon species, but the literature reveals no consensus as to the frequency and ecological circumstances under which chameleon viviparity evolved. We integrated information on reproductive modes for nearly all chameleon species with recently published family-scale phylogenetic and ecological analyses to clarify aspects of reproductive evolution in chameleons. Ancestral-trait reconstructions, after accounting for phylogenetic uncertainty, indicated that viviparity has arisen a minimum of three times in Chamaeleonidae, with each origin of live birth in closed-canopy forests. Our maximum-likelihood optimization therefore did not support the previous hypotheses of one, two or four origins of viviparity in the family. Past claims that arboreality would not allow for evolution of viviparity were also not supported, nor was a recent suggestion that viviparity has reverted to oviparity. However, cold climates of high latitudes and elevations may have selected for viviparity in arboreal chameleons. While peritoneal pigmentation may facilitate viviparity, its role as an exaptation rather than an adaptation remains equivocal without data from a wider range of chameleon species. Based on a comprehensive review of reproductive modes throughout the family, our study has resolved the number of origins of viviparity in Chamaeleonidae and provided evidence that live birth evolved under arboreal conditions on three separate occasions in this enigmatic squamate group. This study also reveals the value of using phylogenetic analysis in a manner that is robust to uncertainty (rather than simple correlational approaches) when the goal is to reconstruct evolutionary sequences and selective pressures.