The evolution of larval morphology and swimming performance in ascidians

The complexity of organismal function challenges our ability to understand the evolution of animal locomotion. To meet this challenge, we used a combination of biomechanics, phylogenetic comparative analyses, and theoretical morphology to examine evolutionary changes in body shape and how those changes affected swimming performance in ascidian larvae. Results of phylogenetic comparative analyses suggest that coloniality evolved at least three times among ascidians and that colonial species have a convergent larval morphology characterized by a large trunk volume and shorter tail length in proportion to the trunk. To explore the functional significance of this evolutionary change, we first verified the accuracy of a mathematical model of swimming biomechanics in a solitary (C. intestinalis) and a colonial (D. occidentalis) species and then ran numerous simulations of the model that varied in tail length and trunk volume. The results of these simulations were used to construct landscapes of speed and cost of transport predictions within a trunk volume/tail length morphospace. Our results suggest that the reduction of proportionate tail length in colonial species resulted in improved energetic economy of swimming. The increase in the size of larvae with the origin of coloniality facilitated faster swimming with negligible energetic cost, but may have required a reduction in adult fecundity. Therefore, the evolution of ascidians appears to be influenced by a trade-off between the fecundity of the adult stage and the swimming performance of larvae.     

Resource allocation in ascidians: reproductive investment vs. other life-history traits

Abstract. To determine patterns of resource allocation among ascidians, we studied 16 colonial and solitary species. We measured investment in reproductive vs. structural material (tunic) both in terms of weight and caloric content, as well as fecundity and degree of larval complexity in the colonial species. Measurements in weight and caloric content were highly correlated in the species studied. A wide range of investment in reproduction was found. Tunic production was related to the growth form of the species, stolonic and solitary species investing less in tunic than massive species, but no significant relationship was found between investment in tunic production vs. reproduction. In colonial species we found that in species with small zooid size, the reproductive investment per zooid was significantly higher. There was a significant negative relationship between investment in reproduction and fecundity. We also found a significant relationship between reproductive investment and larval complexity. The overall trend was that species with low fecundity had large complex larvae and invested the most energy in reproduction.     

Coloniality has evolved once in Stolidobranch Ascidians

Ascidians exhibit a rich array of body plans and life history strategies. Colonial species typically consist of zooids embedded in a common test and brood large, fully developed larvae, while solitary species live singly and usually free-spawn eggs that develop into small, undifferentiated larvae. Ascidians in the order Stolidobranchia include both colonial and solitary species, as well as several species with intermediate morphologies. These include social species, which are colonial but do not live completely embedded in a common test, and a few solitary species that brood embryos and larvae until they are competent to metamorphose. We examined how many times coloniality has evolved within the Stolidobranchia, with phylogenetic analyses using full-length 18S rDNA and partial cytochrome oxidase B sequences for taxa in the families Molgulidae, Styelidae, and Pyuridae. Tunicata orders Phlebobranchia and Stolidobranchia are sister groups, and the family Molgulidae is a monophyletic group and should be raised to the subordinal level, as shown previously by analyses from this lab with partial 18S sequences. In contrast to previous studies, styelids and pyurids are separated into monophyletic groups by ML and Bayesian analyses. We show a single clade within the family Styelidae that contains two colonial (compound) botryllid species, a Symplegma (colonial compound), a colonial (social) species Metandrocarpa taylori, as well as four solitary species, thus confirming that the botryllids are a subfamily of the Styelidae. These results suggest that the ancestor of the Stolidobranchia was solitary and that coloniality has evolved only once within this clade of ascidians. Further phylogenetic analyses of aplousobranch and phlebobranch ascidians will be necessary to understand the number of times that coloniality has evolved within the class Ascidiacea.     

Similarity and diversity in mechanisms of muscle fate induction between ascidian species

Developmental processes can change during evolution at many levels of the ontogeny of an individual. Embryos of solitary ascidians have a largely invariant mode of development, with fixed cleavage patterns and fate maps. Thus the cell lineages and final body plan of the two quite distantly related species considered in this review, Ciona intestinalis and Halocynthia roretzi, are highly similar. However, close comparison of the developmental mechanisms used by these two species provide examples of evolutionary changes and help pinpoint which aspects of development are evolutionarily flexible. Examples of both similarity and diversity are observed in the mechanisms used to generate the full complement of larval muscle. We will describe the changes in muscle-cell lineage, as well as some striking differences in the intercellular signalling pathways used to induce muscle fate. The somewhat surprising conclusion is that in ascidians, as in nematode vulval development, different signalling mechanisms have been adopted to mediate similar interactions between equivalently positioned cells.     

Early activation of adult organ differentiation during delay of metamorphosis in solitary ascidians, and consequences for juvenile growth

Abstract. Many animals have the ability to delay metamorphosis when conditions are unfavorable. This strategy carries obvious benefits, but may also result in severe consequences for lecithotrophic larvae that run low on time and energy. Precocious activation of postlarval developmental programs—so-called anticipatory development—may be adaptive and increase the survival of older, energy-depleted larvae by allowing more rapid metamorphosis. Three of six solitary ascidian species displayed extensive anticipatory development of postlarval structures, similar to heterochronies normally observed for colonial species. The capacity for anticipatory development may be linked to the length of competent period, taxonomic group, or both: members of suborder Phlebobranchia exhibited extensive anticipatory development and long competent periods, but members of suborder Stolidobranchia exhibited little or no anticipatory development and had shorter competent periods. Delay of metamorphosis of up to 3 d did not negatively impact postlarval and juvenile growth rates for any of three species tested, regardless of taxonomic group or length of competent period, although a longer, 7-d, delay resulted in slower postlarval growth in Ciona intestinalis . Anticipatory development of postlarval structures may ameliorate the negative consequences of delay of metamorphosis in C. intestinalis and Ascidiella aspersa , but Molgula socialis showed neither anticipatory development nor a negative impact of metamorphic delay on postlarval fitness. This is the first demonstration that anticipatory development of postlarval structures, normally associated only with colonial ascidians, can occur as a normal part of the development of solitary ascidians.     

Asexual propagation and regeneration in colonial ascidians

Regeneration is widely distributed among the metazoans. However, clear differences exist as to the degree of regenerative capacity: some phyla can only replace missing body parts, whereas others can generate entirely new individuals. Ascidians are animals that possess a remarkable regenerative plasticity and exhibit a great diversity of mechanisms for asexual propagation and survival. They are marine invertebrate members of the subphylum Tunicata and represent modern-day descendants of the chordate ancestor; in their tadpole stage they exhibit a chordate body plan that is resorbed during metamorphosis. Solitary species grow into an adult that can reach several centimeters in length, whereas colonial species grow by asexual propagation, creating a colony of genetically identical individuals. In this review, we present an overview of the biology of colonial ascidians as a paradigm for study in stem cell and regenerative biology. Focusing on botryllid ascidians, we introduce the potential roles played by multipotent epithelia and multipotent/pluripotent stem cells as source of asexual propagation and regenerative plasticity in the different budding mechanisms, and consider the putative mechanism of body repatterning in a non-embryonic scenario. We also discuss the involvement of intra-colony homeostatic processes in regulating budding potential, and the functional link between allorecognition, chimerism, and regenerative potential.     

Vasa expression in a colonial ascidian, Botrylloides violaceus

Evolution of solitary or colonial life histories in tunicates is accompanied by dramatic developmental changes that affect morphology and reproduction. We compared vasa expression in a solitary ascidian and a closely related colonial ascidian, in an effort to uncover developmental mechanisms important during the evolution of these contrasting life histories, including the ability to reproduce by budding. In this study, we explored the origin of germ cells in new buds developing by asexual reproduction in a colonial ascidian, Botrylloides violaceus and compared it to the source of germ cells in a solitary ascidian Boltenia villosa. We studied expression by in situ hybridization of vasa, a DEAD box RNA helicase gene found in germ cells across the metazoans. In B. villosa, bv-vasa mRNA was expressed in putative germ cells and oocytes of adult gonads, and was sequestered into a posterior lineage during embryogenesis. In mature colonies of the ascidian B. violaceus, bot-vasa mRNA was expressed in putative spermatogonia, in oocytes of zooids, and in some circulating cells in the zooids and differentiating buds. We propose that expression of vasa in cells other than gonadal germ cells of zooids in a colonial ascidian may serve as a source of germ-line stem cells in the colony.     

Are the anticipatory pathways in lecithotrophic larvae that delay metamorphosis adaptations? (A review)

During anticipatory development in lecithotrophic larvae that delay metamorphosis, the growth and differentiation of features of the adult action system continue to develop at a slow pace even though they do not become functional. After metamorphosis occurs, the larger size and advanced development of these components may allow juveniles to initially grow at a faster rate than they normally would. Anticipatory development has been demonstrated in archeogastropods, some solitary ascidians and a hydrozoan. In the gastropod Haliotis and the hydrozoan Phialidium anticipatory development increases the initial growth rate of juveniles. In Haliotis and ascidians all of the larvae of a given female that live long enough exhibit anticipatory development. In Phialidium, the ability of a given female to produce larvae that can exhibit anticipatory development is a maternal polymorphic character. In Haliotis and solitary ascidians that exhibit anticipatory development, it appears to be a slower version of the rapid developmental changes that occur in parts of the adult action system at metamorphosis. In Phialidium, developmental changes in relative sizes of the different presumptive regions of the polyp are slowly altered prior to and independently of metamorphosis. Anticipatory development is not linked to the decrease in the size or nutrient reserves of older larvae but to the length of their larval period. From an evolutionary perspective, the mechanisms that operate during anticipatory development are probably of adaptive significance for lecithotrophic larvae of species that spend variable amounts of time in the water column because of a patchy distribution of appropriate settlement cues. The developmental mechanisms that underlie anticipatory development may have been used during the transition from lecithotrophy to planktotrophy.     

The control of the development of a marine benthic community by predation on recruits

Recruitment is an important process in regulating many marine benthic communities and many studies have examined factors controlling the dispersal and distribution of larval immigrants. However, benthic species also have early post-settlement life-stages that are dramatically different from adult and larval stages. Predation on these stages potentially impacts measured recruitment and the benthic populations and communities that ultimately develop.We examined the consequences of post-settlement predation on 1-day-old to 1-month-old recruits of sessile invertebrates at two field sites in southern New England. One site (Breakwater) was in a protected area with few predators and the other (Pine Island) was <1 km away in an open coast area with three different predator guilds: small and large invertebrates and fish. The Breakwater site had been dominated for >10 years by colonial and solitary ascidians. These species were absent from the Pine Island site which was dominated by bryozoans. Our goal was to examine whether post-settlement predation influenced the development and subsequent structure of the epifaunal community.Here we examine long-term changes in community development resulting from post-settlement predation, and contrast these results to those of earlier experiments examining the reductions in observed recruitment by post-settlement predation. Our first long-term experiment examined natural community development at the two sites and whether transplanted communities changed when exposed to the different levels of predation at these sites. The communities that developed at both sites were consistently different from each other and similar to resident communities at their respective sites. On panels transplanted from the Breakwater to Pine Island, solitary ascidians and the colonial ascidian, Botryllus schlosseri, suffered high mortalities on both caged and uncaged treatments, indicative of predation by small predators that could enter cages. Some solitary ascidians did survive inside cages and the colonial ascidian, Botrylloides violaceus, became dominant on all transplanted treatments. On panels transplanted from Pine Island to the Breakwater, ascidians invaded and dominated all treatments except those that were originally caged at Pine Island.In the second long-term experiment, natural communities were allowed to develop on panels exposed at the Breakwater for 1, 2, 3, and 4 weeks. Each set was transplanted to three treatments at Pine Island: open uncaged pilings, caged pilings to exclude fish and large invertebrates, and racks suspended above the bottom to exclude all predators. When 1-week-old communities were transplanted, after 2-3 weeks only bryozoans were found on the open and caged pilings, while colonial ascidians dominated the suspended rack treatment. When older 2-week-old communities were transplanted, colonial ascidians also became dominant in the caged piling treatment and when 3- and 4-week-old communities were transplanted colonial ascidians dominated all three treatments. Solitary ascidians were never abundant on open pilings exposed to fish and large benthic invertebrate predators.Post-settlement predator-prey interactions involved newly settled and juvenile life-stages of a variety of prey species and many invertebrate and vertebrate predator species. The effects of these interactions on recruitment did result in differences in the development and eventual species composition of the communities, even though predators had little if any effect on the adults of the prey species.     

A comparative study of the non-acidic chemically mediated antifoulant properties of three sympatric species of ascidians associated with seagrass habitats

The present study investigated aspects of the antifoulant properties of three sympatric species of ascidians found in seagrass habitats of the Gulf of Mexico, Southern Atlantic Ocean, and Caribbean. Field observations in Saint Joseph Bay, Florida indicate that all three species are common and that the tunic of the solitary ascidian Molgula occidentalis is often heavily fouled, while the outer surfaces of both the colonial ascidians Amaroucium stellatum and Botryllus planus are free of fouling organisms. Antifoulant activities of a suite of increasing hydrophilic organic extracts prepared from the tunic of M. occidentalis and whole colonies of A. stellatum and B. planus were measured using both sympatric microbial (bacteria) and macroinvertebrate (cyprid larvae of Balanus amphitrite) fouling organisms in laboratory bioassays. In addition, field antifoulant assays were conducted by combining organic extracts with controlled-release resin and subsequently coating this material on to acrylic rods deployed in the field for a 72 h period. Extracts of the tunic of M. occidentalis generally did not inhibit bacterial growth. The exception was the methanol extract, which inhibited growth in one of the six marine bacteria tested. Moreover, only the highest concentrations of hexane and methanol tunic extracts tested prevented attachment of cyprid larvae. Field assays revealed no antifoulant activity on rods coated with resin containing extracts of M. occidentalis. Inhibition of both microbial growth and cyprid settlement were much more pronounced in whole-organism extracts of the two colonial ascidians. Most potent were the aqueous methanol extracts of colonies of B. planus and A. stellatum which inhibited growth in five of the six marine bacteria tested. In addition, hydrophilic and lipophilic extracts of the colonial ascidians significantly inhibited attachment of cyprid larvae, in many instances across a wide range of extract concentrations. Field antifoulant assays indicated that extracts of both colonial ascidians inhibited settlement of bryozoans and barnacles. The findings indicate that the colonial ascidians B. planus and A. stellatum possess chemical antifoulant properties. In contrast, the solitary ascidian M. occidentalis appears to either tolerate fouling or possess other non-chemical mechanisms to cope with the risks associated with epibiont overgrowth.     

Larval convergence in a colonial tunicate: the organization of the sarcotubular complex in <Emphasis Type="Italic">Ecteinascidia turbinata</Emphasis> (Perophoridae,Phlebobranchiata, Tunicata,Chordata)

Ecteinascidia turbinata is a colonial ascidian that as an adult shares characters with phlebobranch ascidians, whereas the larvae are similar to aplousobranch ascidian larvae. The sarcotubular complex consists of invaginations of the sarcolemma that contact the sarcoplasmatic reticulum via dyads or triads. If present, the invaginations of the sarcolemma in tunicates have been characterized as laminar or tubular. We comparatively investigated the sarcotubular complex of E. turbinata and seven other tunicate species using 3D-reconstruction techniques based on electron micrographs of serial sections. The mononucleate muscle cells in E. turbinata possess intermediate and close junctions and contain several layers of peripheral myofibrillae. The myofibrillae are surrounded by continuous cisternae of the sarcoplasmic reticulum that forms interconnected rings around the z-bands. The invaginations of the sarcolemma are laminar, contacting the sarcoplasamatic reticulum at the height of the z-bands via dyads and triads. We present a clear definition of character states encountered in Tunicata: laminar invaginations are characterized by a width to length ratio of smaller than 1:20, tubular invagination by a width to length ratio of larger than 1:10. Laminar invaginations are found in stolidobranch ascidians and E. turbinata. Tubular invaginations are present in aplousobranch ascidians and appendicularians. This character state distribution across taxa supports the hypothesis that E. turbinata should be included in Phlebobranchiata as suggested by adult characters and that the larval similarities with Aplousobranchiata arose by convergent evolution. An erratum to this article can be found at     

Ascidian Larvae: The Role of Test Cells in Preventing Hydrophobicity

Abstract Ascidian test cells co-differentiate on the surface of each ovarian oocyte beneath the vitelline coat. They become vacuolated and later occupy the perivitelline compartment of each egg and embryo. In some species, their vacuoles contain submicroscopic granules or filaments called ‘ornaments’ and acidic glycosaminoglycans. These test cells deposit their products on the surface of the larval tunic in late embryogenesis. In these species, the test cells are lost at hatching. In other species, the test cell vacuoles contain acidic glycosaminoglycans, but no ornaments. In these species, the test cells attach to the larval tunic and probably secrete acidic glycosaminoglycans. We deprived the embryos of seven species of ascidians of their test cells and vitelline coats during midembryogenesis. After completing their development, the larvae of both kinds of species were hydrophobic. They easily become trapped on the surface of sea water in cultures. Normal larvae (controls), bearing test cell secretions, are hydrophilic and never become trapped. We infer that negatively charged secretions of the test cells make normal larvae hydrophilic. Some molgulids with direct development have no test cells, no fins and no swimming larva. We reason that the test cells of these species may have been lost during evolution because they no longer had an important role in preventing hydrophobicity.     

Ascidian tunic cells: morphology and functional diversity of free cells outside the epidermis

Abstract. Tunic cells are free cells distributed in the tunic, the integumentary matrix of tunicates. In ascidians, various types of tunic cells have been described both in solitary and in colonial species. Many of them are functionally specialized and are related to the protection of the animal, such as phagocytosis to prevent infection, acid storage to avoid predation, and pigmentation to protect against solar radiation. While some tunic cells are known to play a role in colonial allorecognition, bioluminescence, and algal symbiosis, the functional roles of many cell types still remain to be determined. The composition of tunic-cell types varies among ascidian species, most likely reflecting the functional requirements of the tunic in each species. Although some cell types, e.g., tunic net cells and tunic bladder cells, are restricted to particular taxa of ascidians, tunic phagocytes are found in all known ascidians. Therefore, tunic phagocytes are hypothesized to be basal and shared with ancestral tunicates. In some ascidians, phagocytic cells are involved in other functions, such as pigmentation, intracellular photosymbiosis, and bioluminescence. These specialized phagocytic cells are hypothesized to be derived from tunic phagocytes, suggesting that tunic cells have a high potential to diversify and evolve a wide variety of cellular functions.     

Ascidian neural crest-like cells: phylogenetic distribution, relationship to larval complexity, and pigment cell fate

Migratory neural crest-like cells, which express the cell surface antigen HNK-1 and develop into pigment cells, have recently been identified in the ascidian Ecteinascidia turbinata. Here we use HNK-1 expression as a marker to determine whether neural crest-like cells are responsible for pigment development in diverse ascidian species. We surveyed HNK-1 expression and tyrosinase activity in 12 ascidian species, including those with different adult organizations, developmental modes, and larval sizes and complexities. We observed HNK-1 positive cells in every species, although the timing of HNK-1 expression varied according to the extent of larval complexity. HNK-1 expression was initiated during the late tailbud stage in species in which adult features are formed precociously in large complex larvae. In contrast, HNK-1 positive cells did not appear until the swimming tadpole or juvenile stage in species with small simple larvae in which most adult features appear after metamorphosis. Double labeling experiments indicated that HNK-1 and tyrosinase are expressed in the same subset of pigment-forming mesenchymal cells in species with complex or simple larvae. In addition, the absence of HNK-1 and tyrosinase expression in albino morphs of the colonial ascidian Botryllus schlosseri suggested that the major fate of neural crest-like cells is to become pigment cells. The results suggest that ascidian neural crest-like cells and vertebrate neural crest cells had a common origin during chordate evolution and that their primitive function was to generate body pigmentation.     

Competitive interactions between parasitoid larvae and the evolution of gregarious development

We report experiments using two closely related species of alysiine braconids directed at understanding how gregarious development evolved in one subfamily of parasitoid wasps. Theoretical models predict that once siblicide between parasitoid wasps has evolved, it can only be lost under stringent conditions, making the transition from solitary to gregarious development exiguous. Phylogenetic studies indicate, however, that gregariousness has independently arisen on numerous occasions. New theoretical models have demonstrated that if gregarious development involves reductions in larval mobility, rather than a lack of fighting ability (as in the older models), the evolution of gregariousness is much more likely. We tested the predictions of the older tolerance models (gregariousness based on non-fighting larval phenotypes) and the reduced mobility models (gregariousness based on non-searching larval phenotypes) by observing larval movement and the outcome of interspecific competition between Aphaereta genevensis (solitary) and A. pallipes (gregarious) under multiparasitism. Differences in larval mobility matched the prediction of the reduced mobility model of gregarious development, with the solitary A. genevensis having larvae that are much more mobile. The proportion of hosts producing the solitary species significantly declined after subsequent exposure to females of the gregarious species. This contradicts the prediction of the older models (fighting vs non-fighting phenotypes), under which any competitive interactions between solitary and gregarious larvae will result in a highly asymmetrical outcome, as the solitary species should be competitively superior. The observed outcome of interspecific competition offers evidence, with respect to this subfamily, in favour of the new models (searching vs non-searching phenotypes).     

An evaluation of chemical and physical defenses against fish predation in a suite of seagrass-associated ascidians

The palatability of two solitary and three colonial species of ascidians commonly found in sub-tropical seagrass meadows was evaluated using the abundant, sympatric, omnivorous pinfish Lagodon rhomboides as a model predator. Bite-sized pieces of fresh tissues of both solitary and one of the three colonial ascidian species were unpalatable to fish. Lipophilic and hydrophilic extracts of the three unpalatable species did not cause feeding deterrence indicating that secondary metabolites are not responsible for the lack of palatability. Distaplia bermudensis, the one colonial ascidian that was unpalatable to fish, had a highly acidified outer tunic (pH = 1.5). We tested the ability of acidified agar food pellets (pH = 1.5) to deter pinfish and found that the fish readily ingested acidified pellets. The toughness of the tunic of all five ascidian species was evaluated by measuring the Force (N) required to penetrate the tunic using a penetrometer. Tunic toughness is likely to explain the lack of palatability of the solitary ascidians Styela plicata and Molgula occidentalis as their tunics required a force of > 34 N to penetrate. Tunic toughness may be a particularly effective adaptation for ascidian defense in seagrass habitats where fish with strong crushing jaws, such as those that commonly occur in coral reef systems, are rare.     

Effects of two marinas on the composition of fouling assemblages

The composition of fouling assemblages was surveyed inside and near two fully enclosed marinas using settlement plates. The location of a plate, inside or outside the marina, influenced the abundances of four functional groups of fouling organisms (solitary ascidians, arborescent bryozoans, encrusting bryozoans and colonial ascidians). Transplantation of mature assemblages revealed that reductions in the abundance of arborescent bryozoans inside marinas might be explained by increased growth and recruitment of these bryozoans outside the marina. Surveys of settlement revealed decreased recruitment of encrusting bryozoans inside the marinas, a result consistent with patterns of adult abundance. It is proposed that an increased abundance of solitary ascidians inside marinas may be due to decreased competition. A second survey of adult assemblages was performed with multiple ‘Outside’ sites per marina. Effects of location consistent with the first survey were found for arborescent bryozoans, and in one marina area for solitary ascidians and encrusting bryozoans, but not in the other. Although mechanisms can be proposed to explain the effects of the marina (inside or outside) on the abundances of solitary ascidians, arborescent bryozoans and encrusting bryozoans, the second survey revealed that the effects may vary among marinas.     

The active evolutionary lives of echinoderm larvae

Echinoderms represent a researchable subset of a dynamic larval evolutionary cosmos. Evolution of echinoderm larvae has taken place over widely varying time scales from the origins of larvae of living classes in the early Palaeozoic, approximately 500 million years ago, to recent, rapid and large-scale changes that have occurred within living genera within a span of less than a million years to a few million years. It is these recent evolutionary events that offer a window into processes of larval evolution operating at a micro-evolutionary level of evolution of discrete developmental mechanisms. We review the evolution of the diverse larval forms of living echinoderms to outline the origins of echinoderm larval forms, their diversity among living echinoderms, molecular clocks and rates of larval evolution, and finally current studies on the roles of developmental regulatory mechanisms in the rapid and radical evolutionary changes observed between closely related congeneric species.     

Co-option and dissociation in larval origins and evolution: the sea urchin larval gut

SUMMARY The origin of marine invertebrate larvae has been an area of controversy in developmental evolution for over a century. Here, we address the question of whether a pelagic "larval" or benthic "adult" morphology originated first in metazoan lineages by testing the hypothesis that particular gene co-option patterns will be associated with the origin of feeding, indirect developing larval forms. Empirical evidence bearing on this hypothesis is derivable from gene expression studies of the sea urchin larval gut of two closely related but differently developing congenerics, Heliocidaris tuberculata (feeding indirect-developing larva) and H. erythrogramma (nonfeeding direct developer), given two subsidiary hypotheses. (1) If larval gut gene expression in H. tuberculata was co-opted from an ancestral adult expression pattern, then the gut expression pattern will remain in adult H. erythrogramma despite its direct development. (2) Genes expressed in the larval gut of H. tuberculata will not have a coordinated expression pattern in H. erythrogramma larvae due to loss of a functional gut. Five structural genes expressed in the invaginating archenteron of H. tuberculata during gastrulation exhibit substantially different expression patterns in H. erythrogramma with only one remaining endoderm specific. Expression of these genes in the adult of H. erythrogramma and larval gut of H. tuberculata , but not in H. erythrogramma larval endoderm, supports the hypothesis that they first played roles in the formation of adult structures and were subsequently recruited into larval ontogeny during the origin and evolution of feeding planktotrophic deuterostome larvae.     

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