Reproduction and larval morphology of broadcasting and viviparous species in the Cryptasterina species complex

The Cryptasterina group of asterinid sea stars in Australasia comprises cryptic species with derived life histories. C. pentagona and C. hystera have planktonic and intragonadal larvae, respectively. C. pentagona has the gonochoric, free-spawning mode of reproduction with a planktonic lecithotrophic brachiolaria larva. C. hystera is hermaphroditic with an intragonadal lecithotrophic brachiolaria, and the juveniles emerge through the gonopore. Both species have large lipid-rich buoyant eggs and well-developed brachiolariae. Early juveniles are sustained by maternal nutrients for several weeks while the digestive tract develops. C. hystera was reared in vitro through metamorphosis. Its brachiolariae exhibited the benthic exploration and settlement behavior typical of planktonic larvae, and they attached to the substratum with their brachiolar complex. These behaviors are unlikely to be used in the intragonadal environment. The presence of a buoyant egg and functional brachiolaria larva would not be expected in an intragonadal brooder and indicate the potential for life-history reversal to a planktonic existence. Life-history traits of species in the Cryptasterina group are compared with those of other asterinids in the genus Patiriella with viviparous development. Modifications of life-history traits and pathways associated with evolution of viviparity in the Asterinidae are assessed, and the presence of convergent adaptations and clade-specific features associated with this unusual mode of parental care are examined.     

Evolution of development type in benthic octopuses: holobenthic or pelago-benthic ancestor?

Octopuses of the family Octopodidae are singular among cephalopods in their reproductive behavior, showing two major reproductive strategies: the first is the production of few and large eggs resulting in well-developed benthic hatchlings (holobenthic life history); the second strategy is the production of numerous small eggs resulting in free-swimming planktonic hatchlings (pelago-benthic life history). Here, we utilize a Bayesian-based phylogenetic comparative method using a robust molecular phylogeny of 59 octopus species to reconstruct the ancestral states of development type in benthic octopuses, through the estimation of the most recent common ancestors and the rate of gain and loss in complexity (i.e., planktonic larvae) during the evolution. We found a high probability that a free-swimming hatchling was the ancestral state in benthic octopuses, and a similar rate of gain and loss of planktonic larvae through evolution. These results suggest that in benthic octopuses the holobenthic strategy has evolved from an ancestral pelago-benthic life history. During evolution, the paralarval stage was reduced to well-developed benthic hatchlings, which supports a “larva-first” hypothesis. We propose that the origin of the holobenthic life history in benthic octopuses is associated with colonization of cold and deep sea waters.     

Life‐history predicts past and present population connectivity in two sympatric sea stars

Life‐history traits, especially the mode and duration of larval development, are expected to strongly influence the population connectivity and phylogeography of marine species. Comparative analysis of sympatric, closely related species with differing life histories provides the opportunity to specifically investigate these mechanisms of evolution but have been equivocal in this regard. Here, we sample two sympatric sea stars across the same geographic range in temperate waters of Australia. Using a combination of mitochondrial DNA sequences, nuclear DNA sequences, and microsatellite genotypes, we show that the benthic‐developing sea star, Parvulastra exigua, has lower levels of within‐ and among‐population genetic diversity, more inferred genetic clusters, and higher levels of hierarchical and pairwise population structure than Meridiastra calcar, a species with planktonic development. While both species have populations that have diverged since the middle of the second glacial period of the Pleistocene, most P. exigua populations have origins after the last glacial maxima (LGM), whereas most M. calcar populations diverged long before the LGM. Our results indicate that phylogenetic patterns of these two species are consistent with predicted dispersal abilities; the benthic‐developing P. exigua shows a pattern of extirpation during the LGM with subsequent recolonization, whereas the planktonic‐developing M. calcar shows a pattern of persistence and isolation during the LGM with subsequent post‐Pleistocene introgression.     

Control and Consequences of Alternative Developmental Modes in a Poecilogonous Polychaete

SYNOPSIS. The poecilogonous polychaete Streblospio benedicti(Webster) exhibits both planktotrophic and lecithotrophic modesof larval development. The alternative trophic modes are associatedwith differences in age and size at maturation, offspring number,size and energetic investment, larval planktonic period, morphologyand survivorship. This paper reviews a decade of research intothe control and consequences of the traits associated with planktotrophyand lecithotrophy in S. benedicti. The dominant control on reproductiveand developmental characters is genetic. Significant additivegenetic variance has been detected for egg diameter, fecundity,larval planktonic period and aspects of larval morphology. However,environmental factors such as temperature, food quality andphotoperiod, and intrinsic factors such as maternal age, exertconsiderable influence on non-trophic developmental traits (e.g.,offspring number, size and energy content). Demographic consequencesof development mode are reviewed for field and laboratory demesof S. benedicti dominated by individuals exhibiting either planktotrophyor lecithotrophy. Similar population size structure, fluctuationsin abundance, P: B ratios, and estimated population growth ratesare achieved through trade-offs between survivorship and fecundity. Development mode may best be viewed as a complex set of traitsthat are intimately linked developmentally and evolutionarilyto other aspects of an organism's life history. Greater insightinto the control and consequences of development mode shouldresult from further investigation of these linkages     

Evolution of larval form in the sea star genus Patiriella: conservation and change in the larval nervous system

The organization of the peptidergic system in the larvae of Patiriella species with divergent ontogenies was compared to determine which aspects of neurogenesis are conserved and which are altered in the evolution of development in these sea stars. P. regularis has ancestral-type feeding bipinnaria and brachiolaria larvae and the organization of the nervous system, in association with feeding structures, paralleled the bilateral larval body plan. P. calcar and P. exigua have non-feeding planktonic and benthic brachiolariae, respectively, and there was no trace of the neuronal architecture involved with feeding. The nervous system in the attachment stage brachiolaria was similar in all three species and neuronal organization reflected larval symmetry. Delayed expression of peptidergic lineages to the brachiolaria stage in the lecithotrophs indicates heterochronic change in the timing of neurogenesis or deletion of the ancestral early neurogenic program. The bipinnarial program is suggested to be a developmental module autonomous from the brachiolar one. With a divergence time of less than 10 Ma, the evolution of development in Patiriella has resulted in extensive reduction in the complexity of the larval nervous system in parallel with simplification in larval form. There is, however, strong conservation in the morphology and neuronal architecture of structures involved with settlement.     

Live history evolution in Serpulimorph polychaetes: a phylogenetic analysis

The widely accepted hypothesis of plesiomorphy of planktotrophic, and apomorphy of lecithotrophic, larval development in marine invertebrates has been recently challenged as a result of phylogenetic analyses of various taxa. Here the evolution of planktotrophy and lecithotrophy in Serpulimorph polychaetes (families Serpulidae and Spirorbidae) was studied using a hypothesis of phylogenetic relationships in this group. A phylogenetic (parsimony) analysis of 36 characters (34 morphological, 2 developmental) was performed for 12 selected serpulid and 6 spirorbid species with known reproductive/developmental strategies. Four species of Sabellidae were used in the outgroup. The analysis yielded 4 equally parsimonious trees of 78 steps, with a consistency index (CI) of 0.654 (CI excluding uninformative characters is 0.625). Under the assumption of unweighted parsimony analysis, planktotrophic larvae are apomorphic and non-feeding brooded embryos are plesiomorphic in serpulimorph polychaetes. The estimated polarity of life history transitions may be strengthened by further studies demonstrating an absence of a unidirectional bias in planktotrophy-lecithotrophy transition in polychaetes.     

Strong character incongruence and character choice in phylogeny of sea stars of the Asterinidae

Abstract. Historically, characters from early animal development have been a potentially rich source of phylogenetic information, but many traits associated with the gametes and larval stages of animals with complex life cycles are widely suspected to have evolved frequent convergent similarities. Such convergences will confound true phylogenetic relationships. We compared phylogenetic inferences based on early life history traits with those from mitochondrial DNA sequences for sea stars in the genera Asterina, Cryptasterina , and Patiriella (Valvatida: Asterinidae). Analysis of these two character sets produced phylogenies that shared few clades. We quantified the degree of homoplasy in each character set when mapped onto the phylogeny inferred from the alternative characters. The incongruence between early life history and nucleotide characters implies more homoplasy in the life history character set. We suggest that the early life history traits in this case are most likely to be misleading as phylogenetic characters because simple adaptive models predict convergence in early life histories. We show that adding early life history characters may slightly improve a phylogeny based on nucleotide sequences, but adding nucleotide characters may be critically important to improving inferences from phylogenies based on early life history characters.     

Lipid dynamics in the embryos of Patiriella species (Asteroidea) with divergent modes of development

Evolution of lecithotrophic development in sea stars involved a modification in maternal provisioning from the production of yolk-dominated to lipid-dominated eggs. The dynamics of lipid reserves in the embryos of four Patiriella species differing in their lipid provisions were examined. Patiriella regularis had small yolk protein-dominated eggs (150 microm in diameter) and an ancestral mode of development through planktotrophic larvae. Patiriella calcar, Patiriella exigua and Patiriella pseudoexigua had large eggs (390-440 microm in diameter) and lecithotrophic planktonic, benthic and intragonadal larvae, respectively. Patiriella exigua deposited negatively buoyant eggs containing substantial yolk protein and lipid reserves onto the substratum. In contrast, the planktonic eggs of P. calcar and the intragonadal eggs of P. pseudoexigua were dominated by lipid and were neutrally and positively buoyant, respectively. By the blastula stage there was little trace of lipid in P. regularis embryos. Blastulae of the lecithotrophic developers, by contrast, had conspicuous lipid droplets distributed through their cells. In parallel with the change from cuboidal to columnar epithelium during the blastula to gastrula transition, lipid reserves became redistributed into the basal cytoplasm. The extent of lipid transport reflected the amount of lipid reserves. In P. pseudoexigua embryos with the greatest lipid load, basal shunting was followed by secretion of lipid into the blastocoele where it was stored for the perimetamorphic period. Evolution of lecithotrophy in Patiriella appears to reflect selection to provide metamorphic stages with nutrients normally accrued by feeding larvae with the consequence that early development is burdened by voluminous, potentially inert nutritive stores. Lipid redistribution coincident with a major developmental stage transition may be required to facilitate unimpeded morphogenesis. This phenomenon may be characteristic of lecithotrophic development in echinoderms and appears pre-adaptive for extrusion of lipid in species like P. pseudoexigua with particularly extensive lipid reserves.     

Loss of larval parasitism in parasitengonine mites

Larval Parasitengona are typically parasites, yet at least 29 species of water mites and one species of Trombidiidae forgo larval feeding and any association with a host. Species with non-feeding larvae are isolated cases within species groups or genera where the remaining species have parasitic larvae. Species without larval parasitism occur in at least 14 genera, eight families and four superfamilies of water mites; the loss of larval parasitism is presumably polyphyletic, having occurred at least 21 times. Lineages of water mites with non-feeding larvae frequently exist in parallel with almost identical populations or species that have parasitic larvae. Thus, there is tremendous potential for studies comparing the relative merits of the two life history strategies. Comparisons indicate that adults from lineages with non-parasitic larvae produce smaller numbers of larger eggs; the extra nutrition included in larger eggs permits the larvae to forgo feeding. Non-feeding larvae frequently have wider dorsal plates but reduced leg length, setal length and sclerotization when compared to parasitic larvae from sister lineages. The adults of lineages with non-feeding larvae are frequently smaller in comparison to adults of sister lineages with parasitic larvae. There is no apparent pattern in relation to habitat: lineages lacking larval parasitism occur in streams, temporary ponds and the littoral and planktonic regions of permanent lakes. © Rapid Science Ltd. 1998     

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)     

Why Life Histories Evolve Differently in the Sea

Marine life histories differ from terrestrial life historiesbecause seawater is denser and more viscous than air, becausedesiccation is not a problem for organisms in water, and becausefood is abundant in suspension and solution. (1) Mating andcompetition for paternity in the sea often differs. Female gametesare often spawned freely. Passively dispersed spermatophorescould in some cases provide single paternity to an entire clutchof offspring. Penises of sessile animals reach far for copulation.There are no pollinators. (2) In many clades of benthic marineanimals, greater dispersal of offspring is associated with largeadult size, and greater parental care of offspring and reducedplanktonic larval periods are associated with small adult size.(3) Many benthic marine animals are colonies with modular construction,and these also commonly brood embryos and have short-lived larvae,in contrast to related solitary forms. (4) Unlike dispersalof terrestrial animals, larval dispersal of marine animals isoften obligate with sexual reproduction and often includes aprecompetent period during which larvae cannot settle at goodsites. Unlike terrestrial seeds, marine larvae have no clearadaptations for dispersal, often grow during dispersal, andoften leave bad sites. Feeding planktonic larvae are commonamong marine animals and rare among other aquatic animals, perhapsbecause of persistent aquatic routes between habitable sitesfor marine animals. Peculiarities in marine life histories mayinfluence many aspects of evolution in the sea. Closely relatedsedentary marine animals can differ greatly in larval dispersalwith consequences for recruitment to populations, genetic exchangebetween benthic populations, adaptation to local conditions,sex allocation, interaction with kin, speciation, and extinction.     

A comparative analysis of egg provisioning using mass spectrometry during rapid life history evolution in sea urchins

A dramatic life history switch that has evolved numerous times in marine invertebrates is the transition from planktotrophic (feeding) to lecithotrophic (nonfeeding) larval development—an evolutionary tradeoff with many important developmental and ecological consequences. To attain a more comprehensive understanding of the molecular basis for this switch, we performed untargeted lipidomic and proteomic liquid chromatography‐tandem mass spectrometry on eggs and larvae from three sea urchin species: the lecithotroph Heliocidaris erythrogramma, the closely related planktotroph Heliocidaris tuberculata, and the distantly related planktotroph Lytechinus variegatus. We identify numerous molecular‐level changes possibly associated with the evolution of lecithotrophy in H. erythrogramma. We find the massive lipid stores of H. erythrogramma eggs are largely composed of low‐density, diacylglycerol ether lipids that, contrary to expectations, appear to support postmetamorphic development and survivorship. Rapid premetamorphic development in this species may instead be powered by upregulated carbohydrate metabolism or triacylglycerol metabolism. We also find proteins involved in oxidative stress regulation are upregulated in H. erythrogramma eggs, and apoB‐like lipid transfer proteins may be important for echinoid oogenic nutrient provisioning. These results demonstrate how mass spectrometry can enrich our understanding of life history evolution and organismal diversity by identifying specific molecules associated with distinct life history strategies and prompt new hypotheses about how and why these adaptations evolve.     

Characterization of the lecithotrophic larval development of the temperate New Zealand asterinid Stegnaster inflatus

In the family Asterinidae, development through a planktonic lecithotrophic brachiolaria larva is common and has evolved independently several times. Here, we describe the lecithotrophic development of the asterinid Stegnaster inflatus, a species endemic to New Zealand. Early development through the blastula and gastrula stages is short, with hatching at the brachiolaria stage occurring within 48 hr. After hatching, larvae are negatively buoyant, and without aeration remain near the bottom of the culture containers. The settled benthic juvenile stage was reached in ~2 weeks. The brachiolaria of S. inflatus shares common characteristics with the planktonic brachiolaria of other asterinids in that the brachiolar attachment apparatus comprises three brachia and a central adhesive disc, although the latter is thin and appears to be reduced. Mortensen (1925, Videns kabelige Meddelelser fra Dansk naturhistorisk Forening i København, 79 (15), 261‐420) had hypothesized that individuals of S. inflatus might brood within the “cave” formed in the interambulacral space between the arms. We found no evidence for brooding, but hypothesize that S. inflatus may have demersal development, on or near the bottom, which has implications for larval dispersal and population structure.     

Distribution of echinoderms and their larvae in the southwestern Peter the Great Bay, Sea of Japan

Based on results of processing planktonic and benthic samples collected in 1996 and 1997, a spatial distribution of echinoderms in the bottom and of their larvae in the plankton were collated for the water area of the Southern region of the Far Eastern State Biosphere Marine Reserve. Some correlation between distributions of the adult and larval sea star Asterias amurensis in July was revealed. At the same time, there was no correlation between distributions of larvae and adult individuals of the brittle star Ophiura sarsi and sea urchin Echinocardium cordatum, which are most abundant in the area. The size structure of bottom populations of the brittle stars O. sarsi and Amphiodia fissa in the studied area was assessed. The correlation coefficient between the distribution of young-of-the-year and the population density was 0.47 in O. sarsi and 0.74 in A. fissa respectively, which implied a selective settling of larvae of those species in areas inhabited by adult brittle stars. Recruitment of bottom populations from 1996 spawning was 5% in O. sarsi and 3.3% in A. fissa.     

Lectin histochemistry of the hyaline layer around the larvae of Patiriella species (Asteroidea) with different developmental modes.

Larvae of sea stars are surrounded by an extracellular matrix called the hyaline layer. The lectin-binding properties of this matrix were investigated in an ultrastructural study of Patiriella species having different modes of development. The planktonic bipinnaria and brachiolaria of P. regularis and the planktonic brachiolaria of P. calcar demonstrated the same labeling of the hyaline layer for three lectins: Con A, SBA, and WGA. In both species the outer coarse meshwork stained for all three lectins, whereas the intervillous layer displayed patchy labeling. In the benthic brachiolaria of P. exigua, the outer coarse meshwork displayed heavy labeling for all three lectins. The heavy labeling of the outer coarse meshwork of P. exigua compared with that of the other species suggests an increased number of lectin binding sites in the hyaline layer of this species. The similar ultrastructure and histochemistry of the hyaline layer of P. regularis and P. calcar may reflect similar requirements of their extracellular cover in their planktonic environment. Lectin labeling shows that hypertrophy of the hyaline layer of P. exigua, in particular the outer coarse meshwork, involves elaboration of the carbohydrate composition of the matrix. Modifications seen in the ultrastructure and histochemistry of the hyaline layer of P. exigua appear to be associated with the evolution of benthic development.     

A new poecilogonous species of sea slug (Opisthobranchia: Sacoglossa) from California: comparison with the planktotrophic congener Alderia modesta (Loven, 1844)

Cryptic species are increasingly recognized as commonplace amongmarine gastropods, especially in taxa such as shell-less opisthobranchsthat lack many discrete taxonomic characters. Most cases ofpoecilogony, the presence of variable larval development withina single species, have historically turned out to representcryptic species, with each possessing a single canalized typeof development. One well-characterized example of poecilogonywas attributed to the sacoglossan opisthobranch Alderia modesta;in southern California, slugs resembling this member of a monotypicgenus produce both long-lived, planktotrophic and short-lived,lecithotrophic larvae. Paradoxically, however, A. modesta isexclusively planktotrophic everywhere else in the northern Pacificand Atlantic Oceans. A recently completed molecular study foundthat slugs from poecilogonous populations south of Bodega Harbor,California, comprise an evolutionarily distinct lineage separatefrom northern, strictly planktotrophic slugs. We now describethe southern species as A. willowi n. sp., based on differencesin morphology of the dorsum and radula, characteristics of theegg mass, larval development mode and nuclear and mitochondrialgenetic markers. A DNA barcode is provided, based on 27 fixeddifferences in the cytochrome c oxidase subunit I gene thatcan reliably differentiate Pacific specimens of Alderia species.Genetic and morphological data are concordant with developmentalevidence, confirming that A. willowi is a true case of poecilogony.An improved understanding of the ecological differences betweenthese sister taxa may shed light on the selective pressuresthat drove the evolution of lecithotrophy in the southern species. (Received 1 November 2005; accepted 20 September 2006)     

Abbreviation of larval development and extension of brood care as key features of the evolution of freshwater Decapoda

The transition from marine to freshwater habitats is one of the major steps in the evolution of life. In the decapod crustaceans, four groups have colonized fresh water at different geological times since the Triassic, the freshwater shrimps, freshwater crayfish, freshwater crabs and freshwater anomurans. Some families have even colonized terrestrial habitats via the freshwater route or directly via the sea shore. Since none of these taxa has ever reinvaded its environment of origin the Decapoda appear particularly suitable to investigate life‐history adaptations to fresh water. Evolutionary comparison of marine, freshwater and terrestrial decapods suggests that the reduction of egg number, abbreviation of larval development, extension of brood care and lecithotrophy of the first posthatching life stages are key adaptations to fresh water. Marine decapods usually have high numbers of small eggs and develop through a prolonged planktonic larval cycle, whereas the production of small numbers of large eggs, direct development and extended brood care until the juvenile stage is the rule in freshwater crayfish, primary freshwater crabs and aeglid anomurans. The amphidromous freshwater shrimp and freshwater crab species and all terrestrial decapods that invaded land via the sea shore have retained ocean‐type planktonic development. Abbreviation of larval development and extension of brood care are interpreted as adaptations to the particularly strong variations of hydrodynamic parameters, physico‐chemical factors and phytoplankton availability in freshwater habitats. These life‐history changes increase fitness of the offspring and are obviously favoured by natural selection, explaining their multiple origins in fresh water. There is no evidence for their early evolution in the marine ancestors of the extant freshwater groups and a preadaptive role for the conquest of fresh water. The costs of the shift from relative r‐ to K‐strategy in freshwater decapods are traded‐off against fecundity, future reproduction and growth of females and perhaps against size of species but not against longevity of species. Direct development and extension of brood care is associated with the reduction of dispersal and gene flow among populations, which may explain the high degree of speciation and endemism in directly developing freshwater decapods. Direct development and extended brood care also favour the evolution of social systems, which in freshwater decapods range from simple subsocial organization to eusociality. Hermaphroditism and parthenogenesis, which have evolved in some terrestrial crayfish burrowers and invasive open water crayfish, respectively, may enable populations to adapt to restrictive or new environments by spatio‐temporal alteration of their socio‐ecological characteristics. Under conditions of rapid habitat loss, environmental pollution and global warming, the reduced dispersal ability of direct developers may turn into a severe disadvantage, posing a higher threat of extinction to freshwater crayfish, primary freshwater crabs, aeglids and landlocked freshwater shrimps as compared to amphidromous freshwater shrimps and secondary freshwater crabs.     

Effects of temporary food limitation on survival and development of brachyuran crab larvae

As a consequence of the combined effects of prey patchinessand diel or tidal vertical migrations in the water column, decapodcrustacean larvae may experience temporal or spatial variabilityin the availability of planktonic food. In a laboratory study,we evaluated effects of temporarily limited access to prey onthe larvae of three species of brachyuran crabs, Chasmagnathusgranulata, Cancer pagurus and Carcinus maenas. Stage-I zoeaewere fed ad libitum for 4 or 6 h per day (20 or 25% treatments;6 h tested in C. pagurus only), and rates of larval survivaland development were compared with those observed in continuouslyfed control groups (24 h, 100%). In C. granulata, we also testedif intraspecific variability in initial biomass of freshly hatchedlarvae originating from different broods has an influence onearly larval tolerance of food limitation. Moreover, we exposedembryos and larvae of this estuarine species to moderately decreasedsalinities to identify possible interactions of osmotic andnutritional stress. Finally, we evaluated in this species theeffect of food limitation on survival from hatching throughall larval instars to metamorphosis. In all three species, limitedaccess to prey had only weak or insignificant negative effectson survival through the Zoea-I stage. The strength of the effectsof temporary food limitation varied in C. granulata significantlyamong broods. However, no significant relationships were foundbetween initial larval biomass (C content) and either survivalor development duration. Strongly decreased survival to metamorphosiswas found when food limitation continued throughout larval development.Thus, early brachyuran crab larvae are well adapted to transitorylack of planktonic food. The capability of the Zoea-I stageof C. granulata to withstand nutritional stress also under conditionsof concomitant salinity stress allows them to exploit variousbrackish environments within estuarine gradients. However, continuedexposure to limited access to planktonic prey may exceed thenutritional flexibility of C. granulata larvae.     

Phylogenetic analyses of mode of larval development

Phylogenies based on morphological or molecular characters have been used to provide an evolutionary context for analysis of larval evolution. Studies of gastropods, bivalves, tunicates, sea stars, sea urchins, and polychaetes have revealed massive parallel evolution of similar larval forms. Some of these studies were designed to test, and have rejected, the species selection hypothesis for evolutionary trends in the frequency of derived larvae or life history traits. However, the lack of well supported models of larval character evolution leave some doubt about the quality of inferences of larval evolution from phylogenies of living taxa. Better models based on maximum likelihood methods and known prior probabilities of larval character state changes will improve our understanding of the history of larval evolution.