Evolution of the molluscan body plan: the case of the anterior adductor muscle of bivalves

The separated shell plates with the rearranged musculature (adductor muscle) is a novelty for bivalves. Despite its importance in the bivalve bodyplan, the development of the anterior adductor muscle remains unresolved. In this study, we investigate the myogenesis of the bivalve species Septifer virgatus to reveal the developmental origin of the larval muscles in bivalves, focusing on the anterior adductor muscle. We observed that larval retractor muscles are differentiated from the ectomesoderm in bivalves, and that the anterior adductor muscles are derived from primordial larval retractor muscles via segregation of the myoblast during the veliger larval stage. Through the comparative study of myogenesis in bivalves and its related taxa, gastropods, we found that both species possess myoblasts that emerge bilaterally and later meet dorsally. We hypothesize that these myoblasts, which are a major component of the main larval retractor in limpets, are homologous to the anterior adductor muscle in bivalves. These observations imply that the anterior adductor muscle of bivalves evolved as a novel muscle by modifying the attachment sites of an existing muscle.     

Diet composition of two larval headwater stream salamanders and spatial distribution of prey

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First record of Oxytorinae (Hymenoptera: Ichneumonidae) from South America,with description of a new species of <Emphasis Type="Italic">Oxytorus</Emphasis> Förster, 1869

The monotypic subfamily Oxytorinae is recorded for the first time from South America. Oxytorus peruvianus sp. n. from a Peruvian premontane rain forest is described and illustrated. Information on the habitat and phenology of the new species is reported.     

Mutualism between Thisbe irenea butterflies and ants, and the role of ant ecology in the evolution of larval-ant associations

A facultative mutualism between the riodinid butterfly Thisbe irenea and the ponerine ant Ectatomma ruidum is described from Panama. Ants protect larvae against attacks of predatory wasps, but not against tachinid parasitoids. Several potential sources of ecological variation affecting the larval survival of Thisbe irenea are noted. A preliminary means of testing the ability of larvae to appease ants is described that may be applied to all butterfly-ant systems. Observations and literature records indicate that ant taxa which tend butterfly larvae are the same taxa that tend extrafloral nectaries and Homoptera. A general hypothesis for the evolution of myrmecophily among butterflies suggests that ant taxa which utilize secretions in their diet are major selective agents for the evolution of the larval ant-organs, and hence, ant-larval mutualisms. This idea is extended to suggest how appeasement of predaceous ant taxa through the use of larval ant-organs can influence an ant-larval relationship, eventually leading to mutualism.     

A preliminary distributional study of fish larvae near a ribbon coral reef in the Great Barrier Reef

Fish larvae from horizontal plankton tows along a single transect near outer ribbon reefs of the Great Barrier Reef in spring 1979 and summer 1980 had persistent distributional patterns. Larvae were identified to family and divided into young (preflexion) and old (postflexion) larvae, thus giving 28 taxa abundant enough for analysis. Non-uniform larval distributions were found for 81% of the 16 reef fish taxa with non-pelagic eggs, but for only 17% of the six reef fish taxa with pelagic eggs. Most differences in larval concentration were between the lagoonal and seaward sides of the reef. Only tripterygiid larvae had highest concentration just seaward of the reef, while larvae of 12 reef and three oceanic fish taxa occurred in highest concentrations on the lagoonal side of the reef. In five taxa of reef fishes, higher larval concentrations were found in the lagoonal backreef compared with the mid-lagoon habitat; but the reverse was not found in any taxon. Eleven taxa had indeterminate distributions, (i.e. no difference in concentration between stations). Mechanisms responsible for the distribution remain unknown, but we suggest that the view which considers fish larvae to be passively-drifting particles is unjustified without more information on larval behaviour.     

Better safe than sorry? A Fabaceae species exhibits unfavourable pollen properties for developing bee larvae despite its hidden anthers

Empirical evidence suggests that pollen chemistry plays an important role in shaping the pollen host spectra of many bee species. Although the underlying mechanisms are poorly understood, pollen diets of several plant taxa have experimentally been found to impede larval development of unspecialized bees. The pollen of all plant taxa, for which such a detrimental effect on bee larval development has been observed so far, is freely accessible in the flowers and thus easily harvestable for flower visitors, suggesting that this pollen might be chemically protected in order to reduce its loss to pollen-feeding animals. In the present study, we compared larval performance of five solitary bee species on pollen diets of the two Fabaceae species Onobrychis viciifolia and Lotus corniculatus, which have their anthers concealed inside the flowers, with that on control diets composed of host pollen provisions. As the complex flower morphology of the two Fabaceae species already considerably narrows the spectrum of pollen harvesting bee taxa, which might supersede costly chemical protection of the pollen, we expected bees that usually do not exploit Fabaceae to develop well on Onobrychis and Lotus pollen diets. Larval survival on the Onobrychis pollen diet was successful for all five bee species tested. In contrast, larval survival on the Lotus pollen diet was reduced in three species despite the fact that Lotus flowers are more difficult to exploit for pollen than Onobrychis flowers. We conclude that there is no trade-off between pollen concealment and pollen defence in Lotus and that pollen of morphologically complex flowers with a restricted visitor spectrum is not necessarily an easy-to-use nutritional source.     

Evolution of larval size in cyclostome bryozoans

Cyclostomes are the only order of stenolaemate bryozoans living today. The non-feeding larvae of modern cyclostomes metamorphose on settlement to produce a calcified dome-shaped protoecium. Protoecial diameter provides a proxy for larval size. The sparse data available on living cyclostomes suggests that protoecial diameter is about one-and-a-half times greater than larval width. Here we use protoecial diameter to estimate larval sizes in fossil and Recent cyclostome species. A total of 233 protoecia were measured, 143 from Recent cyclostomes and 90 from fossil cyclostomes, of which 84 came from the Jurassic. Protoecial diameter ranged from 82.5 to 690 μm, with 89% of protoecia having diameters between 100 and 300 μm. A comparison of 30 Jurassic with 51 Recent taxa of tubuliporine cyclostomes showed a significant difference in size frequency. Although the Recent taxa have a larger size range (83–465 μm) than the Jurassic taxa (125–249 μm), Recent species have a lower mode (125–150 μm) than the Jurassic species (175–200 μm). Most Jurassic cyclostomes may therefore have had larger larvae than their extant relatives. Reduction in larval size may be a component of the previously hypothesized reduction in overall body size resulting from competitive displacement by cheilostome bryozoans.     

Chironomid biomass determination from larval shape

SUMMARY.

• 1 Ash-free dry weights and dry weights of each larval instar were determined for nine chironomid taxa using unpreserved material. Larval weight (wμg) was related to body length measurements (l mm) by the equation In w= In a+b In l.
• 2 For a given taxon. the constants In a and b were dependent on larval size and shape, respectively: b was found to increase with larval length, while taxa with characteristically stout larvae yielded higher values for In a.
• 3 Comparison of measurements of freshly-killed chironomid larvae with measurements taken after preservation in 70% ethanol showed no change in body shape due to preservation. Length measurements from larvae preserved in this way are therefore suitable for use in the calculation of larval biomass when the relationship to the unpreserved weight is known.
• 4 Maximum larval lengths and mean body proportions (length: width ratios) are reported for thirty-two chironomid taxa.

     

Body size,trophic level,and the use of fish as transmission routes by parasites

Within food webs, trophically transmitted helminth parasites use predator–prey links for their own transfer from intermediate prey hosts, in which they occur as larval or juvenile stages, to predatory definitive hosts, in which they reach maturity. In large taxa that can be used as intermediate and/or definitive hosts, such as fish, a host species’ position within a trophic network should determine whether its parasite fauna consists mostly of adult or larval helminths, since vulnerability to predation determines an animal’s role in predator–prey links. Using a large database on the helminth parasites of 303 fish species, we tested whether the proportion of parasite species in a host that occur as larval or juvenile stages is best explained by their trophic level or by their body size. Independent of fish phylogeny or habitat, only fish body length emerged as a significant predictor of the proportion of parasites in a host that occur as larval stages from our multivariate analyses. On average, the proportion of larval helminth taxa in fish shorter than 20 cm was twice as high as that for fish over 100 cm in length. This is consistent with the prediction that small fishes, being more vulnerable to predation, make better hosts for larval parasites. However, trophic level and body length are strongly correlated among fish species, and they may have separate though confounded effects on the parasite fauna exploiting a given species. Helminths show varying levels of host specificity toward their intermediate host when the latter is the downstream host involved in trophic transmission toward an upstream definitive host. Given this broad physiological compatibility of many helminths with fish hosts, our results indicate that fish body length, as a proxy for vulnerability to predators, is a better predictor of their use by helminth larvae than their trophic level based on diet content.     

Leaf-Associated Bacterial and Fungal Taxa Shifts in Response to Larvae of the Tree Hole Mosquito, <Emphasis Type="Italic">Ochlerotatus triseriatus</Emphasis>

Larvae of the eastern tree hole mosquito, Ochlerotatus triseriatus (Say), and related container-breeding species are known to feed upon substrate-associated microorganisms. Although the importance of these microbial resources to larval growth has been established, almost nothing is known about the taxonomic composition and dynamics of these critical microbial food sources. We examined bacterial and fungal community compositional changes on oak leaves tethered in natural tree hole habitats of O. triseriatus. We eliminated larvae experimentally in a subset of the tree holes and examined 16S rDNA gene sequences for bacteria and ergosterol concentrations and 18S rRNA gene sequences for fungi collected from leaf material subsamples. Leaf ergosterol content varied significantly with time, but not treatment. Principal component analysis (PCA) was used to compare microbial taxonomic patterns found in leaves incubated with or without larvae present, and we found that larval presence affected both bacterial and fungal groups, either from loosely attached or strongly adherent categories. Bacterial communities generally grouped more tightly when larvae were present, and class level taxa proportions changed when larvae were present, suggesting selection by larval feeding or activities for particular taxa such as members of the Bacteroidetes, Alphaproteobacteria, and Betaproteobacteria classes. Fungal taxa composite scores also separated along PC axes related to the presence of larvae and indicated larval feeding effects on several higher taxonomic groups, including Saccharomycetes, Dothideomycetes, and Chytridiomycota. These results support the hypothesis that larval mosquito feeding and activities altered microbial communities associated with substrate surfaces, potentially leading to decreased food value of the resource and affecting decomposition of particulate matter in the system.     

Larval morphology and biology of oxycorynine weevils and the higher phylogeny of Belidae (Coleoptera, Curculionoidea)

Phylogenetic relationships among members of the family Belidae (Curculionoidea) were reconstructed through cladistic analysis using 58 characters and 17 terminals. The characters were from larval morphology (30), adult morphology (25) and biology regarding larval host-plants and feeding habits (three). They were scored for exemplar taxa in 17 genera, representing different belid subfamilies and tribes, plus two outgroup taxa in Megalopodidae and Nemonychidae. The sampled genera included all those for which larval and adult information is available, and two known only from adults. New information on the larvae and biology of two oxycorynines is provided. These are the Chilean Oxycraspedus cribricollis , whose larvae live in decayed female strobili of the gymnosperm Araucaria araucana , and Hydnorobius hydnorae from Argentina, whose larvae, described and illustrated in the present paper, develop inside the flower and fruit bodies of Prosopanche americana (Hydnoraceae), a root-parasitic angiosperm. The relationships proposed by the single optimal cladogram resulting from simultaneous analysis of all taxa and characters are recovered by one of three optimal cladograms based on the larval data set alone. The cladogram justifies a revised classification of Belidae in two sister subfamilies: Belinae (with tribes Pachyurini, Agnesiotidini and Belini) and Oxycoryninae (with tribes Oxycorynini and Aglycyderini). It summarizes larval and adult synapomorphies defining the family Belidae, subfamilies and tribes. Based on the phylogenetic tree, the evolution of biological traits is traced. Larval development in vegetative organs of conifers is ancestral in Belidae. A shift to reproductive structures characterizes the Oxycorynini, a habit which was conserved while several shifts to distantly related host-plant groups occurred.     

Larval Case Architecture and Implications of Host‐Plant Associations for North American Coleophora (Lepidoptera; Coleophoridae)1

The case‐bearing moths of North America are represented by a single genus, Coleophora, which contains approximately 144 described species. All are external seed or leaf miners that inhabit portable silk cases during most of the larval stage. Architectural and ecological characters from larval cases were used in cladistic analysis to investigate existing case groups for 32 North American species of Coleophora. Cladistic analysis confirmed monophyly of certain case groups, but not of others. Host‐plant preferences were also examined. The pattern of host plant use reflects more closely preference for certain plant tissues (seeds versus leaves) and growth forms (herbaceous versus woody) with exploitation of different plant taxa, rather than preference for certain plant taxa with exploitation of different plant tissues.     

Larval and early post-larval shell of three deep-sea Scaphopoda (Mollusca) from the southwest Atlantic

The larval shells of Antalis circumcincta (Watson, 1879) (order Dentaliida), Pertusiconcha callithrix (Dall, 1889) (order Gadilida) and of an undetermined species of uncertain systematic position are described. The material studied comprises mainly samples from deep waters, collected by expeditions along the southeast coast of Brazil. The larval shell of the three taxa matches other types previously described in the literature. Antalis circumcincta and P. callithrix have four regions (protoconch A, protoconch B, teleoconch A, teleoconch B), but differ in dimensions and sculpture from each other, while the undetermined species has three regions (protoconch A, protoconch B, teleoconch B). A morphometric approach combined with a discriminant analysis also indicates that the three taxa are significantly distinct. This study confirms patterns of larval shells at the taxonomic rank of orders but other supraspecific patterns remain uncertain.     

COMPOSITION,ABUNDANCE AND SEASONALITY OF LARVAL FISH IN RICHARDS BAY HARBOUR,KWAZULU-NATAL,SOUTH AFRICA

Summary

The present study examined the composition, abundance and seasonality of fish larvae occurring in Richards Bay Harbour, over an 18-month period from June 1991 to November 1992. Ichthyoplankton samples were collected every six weeks at top, mid and bottom depths in the dredged channel 100 to 200 m from the harbour entrance. A total of 7 163 fish larvae, representing 105 taxa and 53 families, was collected throughout the study period. The dominant families were the Engraulidae and Gobiidae representing 50% and 37% of the total catch, respectively. The most abundant larvae were those of the thorny anchovy, Stolephorus holodon, and an unidentified goby, Gobiid 12 (32% and 30% of the total catch, respectively). Estuarine-associated taxa dominated the total catch in terms of density (61%), with 11 out of the 28 taxa being totally dependent on estuaries at some stage in their life cycle. Turbidity and temperature were significant variables accounting for 27% of the variation in larval densities of estuarine-dependent species. The main peak in larval density occurred in January 1992 (summer) with a mean larval density of 160 larvae per 100 m³. Larvae of estuarine-associated taxa were predominantly at the flexion and postflexion developmental stages. Since larval densities of certain estuarine-associated species were significantly greater in bottom samples and on flood tides, it is suggested that selective tidal stream transport is a recruitment mechanism used by these species. The present study concluded that it is important to maintain and preserve the estuarine component of Richards Bay Harbour so that the system can function as an estuarine nursery for many of the fish species depending on it at certain stages in their life cycle.     

Nectaropolliniferous Sources Used by Trigona (Tetragonisca) Angustula in Chiapas,Southern México

The nectaropolliniferous flora used by Trigona (Tetragonisca) angustula (Apidae, Meliponinae), was studied by melissopalinological analysis at Chiapas, southern Mexico, in two zones with different altitudes and vegetation (Santa Teresita and Unión Juárez). In both areas monthly samples of honey, pollen, and larval feed were taking for a full year, from a nest established expressly for the purpose. A total of 105 plant species were identified at Santa Teresita, of which only 28 taxa were present in percentages higher than 10; of these, seven taxa are nectariferous, ten nectaropolliniferous and eleven polliniferous sources, of which four taxa are found exclusively in larval food. The values of niche size (H') at Santa Teresita indicated less diversity and availability of sources, in comparison with Unión Juárez, where the evenness values (J') have a tendency toward heterogeneity. At Unión Juárez, a total of 127 plant species were registered of which only 30 taxa were present in frequencies higher than 10%; of these ten species are nectariferous, five taxa are nectaropolliniferous, and 15 species are polliniferous, and of the last category three taxa are found exclusively in brood cells. At Unión Juárez the values of niche size (H') for the three sources are higher than at Santa Teresita, and the evenness (J') values are homogeneous, and showed tendency toward evenness, indicating more diversity and availability in this area.     

Identification of asteroid genera with species capable of larval cloning

Asexual reproduction in larvae, larval cloning, is a recently recognized component of the complex life histories of asteroids. We compare DNA sequences of mitochondrial tRNA genes (Ala, Leu, Asn, Pro, and Gln) from larvae in the process of cloning collected in the field with sequences from adults of known species in order to identify asteroid taxa capable of cloning. Neighbor-joining analysis identified four distinct groups of larvae, each having no, or very little, sequence divergence (p distances ranging from 0.00000 to 0.02589); thus, we conclude that each larval group most likely represents a single species. These field-collected larvae cannot be identified to species with certainty, but the close assemblage of known taxa with the four larval groups indicates generic or familial identity. We can assign two of the larval groups discerned here to the genera Luidia and Oreaster and another two to the family Ophidiasteridae. This study is the first to identify field-collected cloning asteroid larvae, and provides evidence that larval cloning is phylogenetically widespread within the Asteroidea. Additionally, we note that cloning occurs regularly and in multiple ways within species that are capable of cloning, emphasizing the need for further investigation of the role of larval cloning in the ecology and evolution of asteroids.     

TRAIT MAPPING AND SALIENCE IN THE EVOLUTION OF EUSOCIAL VESPID WASPS

The multiple independent origins of eusociality in the insect order Hymenoptera are clustered in only four of more than 80 families, and those four families are two pairs of closely related taxa in a single part of the order. Therefore, although ordinal-level characteristics can contribute to hymenopteran eusocial evolution, more important roles have been played by traits of infraordinal taxa that contain the eusocial forms. Many factors have been proposed and discussed, but assessments of traits' salience to eusocial evolution have heretofore not been joined to phylogenetics. In the present analysis, cladograms of superfamilies and families of Hymenoptera and of the family Vespidae are used to ordinate the appearance of traits that play roles in vespid eusociality. Proximity of traits' first appearance to the origin of eusocial Vespidae is taken as one measure of traits' salience to vespid eusocial evolution. Traits that subtend only eusocial taxa and that are uniquely associated with eusociality have foundations in more general traits that subtend more inclusive taxa. No single trait is uniquely causative of vespid eusocial evolution. High-salience traits that closely subtend vespid eusociality include nesting, oviposition into an empty nest cell, progressive provisioning of larvae, adult nourishment during larval provision malaxation, and inequitable food distribution among nestmates. The threshold characteristic of Polistes-grade eusociality is life-long alloparental brood care by first female offspring who remain, uninseminated, at their natal nest. Traits directly associated with occurrence of such workers are larva-adult trophallaxis, which can foster relatively low larval nourishment early in a colony cycle, and protogyny and direct larval development, which combine to yield restricted mating opportunities for female offspring that are the first to emerge in the colony cycle. Trait mapping suggests no role for asymmetry of relatedness due to haplodiploidy, but it suggests high salience for haplodiploidy as a mechanism enabling the production of all-female clutches of first offspring.     

The transition from planktotrophy to lecithotrophy in larvae of Lower Palaeozoic Rhynchonelliform brachiopods

Criteria are established for defining the presence of protegula formed on embryonic or larval mantle in representative genera of Lower Palaeozoic Obolellata, Strophomenata and Rhynchonellata. Width was used to define protegular type. Taxa with only an embryonic protegulum are inferred to have had lecithotrophic larvae while taxa with a larval protegulum or an embryonic protegulum surrounded by a larval protegulum are inferred to have had planktotrophic larvae. All or most of the taxa examined in the Obolellata, the Strophomenata and the orders Protorothida and Orthida in the Rhynchonellata had planktotrophic larvae. In the Pentamerida a minority of genera had only a larval, or an embryonic and a larval protegulum while a majority had protegular widths indicating lecithotrophy. In the orders Rhynchonellida, Atrypida, Athyrida and Spiriferida derived from the Pentamerida (with the exception of one species in the Atrypida) a number of the genera had protegular widths indicating lecithotrophy. It is suggested that the onset of lecithotrophy in the Pentamerida was associated with a developmental innovation in which the mantle lobe of the larva was reflected over the apical lobe during the process of metamorphosis. This evolutionary innovation probably occurred during the late Cambrian or early Ordovician and was subsequently inherited during the process of cladogenesis.     

Temporal and spatial patterns in ichthyoplankton assemblages in bay and open coastal environments

The larval fish fauna occurring in temperate bay and shelf waters off Victoria, southern Australia, was found to be diverse, comprising taxa from 52 families. The most abundant groups collected were gobiids, tripterygiids, gobiesocids and clupeids. Fish egg concentrations were highest during spring and summer (September to February). Eggs of the Australian anchovy Engraulis australis occurred mainly during spring (September to November). Total larval fish concentrations were highest during summer (December to February), and were significantly higher at 1 km than 2 and 5 km from shore in offshore samples. Larval concentrations of a number of families, mainly reef‐associated taxa that attach their eggs to hard substrata, were also higher nearer to shore. These larvae are more developed upon hatching than those of pelagic spawners and more capable of avoiding passive drift. Multivariate analyses found that larval taxonomic composition did not vary significantly with distance from shore, but that seasonal and monthly groupings were evident, with different taxa dominating at different times of the year. Larvae of the families Gobiidae and Tripterygiidae occurred in all months, but were less abundant during winter. Spatial differences in the larval fish assemblage between offshore samples and samples taken in the bay were only apparent during summer. This was primarily due to a higher abundance of seagrass‐associated species, such as syngnathids and hemiramphids, utilizing specific habitats in the bay.