Progenesis as a principle in meiofauna evolution

The interstitial meiofaunal taxa may have evolved in very different ways, which are shortly discussed and schematically represented. Several taxa reveal the possibility of an evolutionary pathway by early maturation of larvae or juvenile stages of large size species with a pelago-benthic life-cycle. This presumed progenetic (= neotenic) evolution can be sustantiated in, for example, various Polychaeta and Crustacea by larval or juvenile morphological features. The progenetic origin of several of these taxa is also strongly supported by morphological series, of which species conforming well with typical larvae form the endpoint. The monophyletic origin of similar progenetic taxa, however, is only proved with difficulties. The theoretical available phylogenetic reconstructions for a series of progenetic taxa are discussed. It is emphasized that progenetic evolution is not regressive or degenerate but novel morphological features like derived reproductive organs are characteristic indications of the progenetic origin of a species. In general, progenesis may have a significant role in the evolution of higher taxa in the interstitial habitat.     

Disentangling prey-handling efficiency of larval newts through multivariate prey trait analysis

Multivariate prey trait analysis is a functional approach to understand predator–prey relationships. In the present study, six prey macroinvertebrate ecological traits were used to identify several key factors in the handling efficiency of seven predatory larval newt species. The results revealed a remarkable similarity in the feeding behaviour among species, suggesting a foraging behaviour convergence among species in the prey-handling efficiency of larval newts. Nevertheless, larvae of some newt species showed clear preferences for particular categories of ecological trait; for example, Lissotriton vulgaris tended to feed on macroinvertebrate taxa with random trajectory, and Calotriton asper fed on macroinvertebrate taxa with high agility. In contrast, Mesotriton alpestris showed a clear ability to feed on prey with movable accessories (cases/tubes) and patterned concealment. This study shows how multivariate approaches can complement traditional diet analyses, and the method has wide applicability across life-stages and species.     

Bionomics,behaviour and immature stages of Pelecotoma fennica (Paykull) (Coleoptera: Rhipiphoridae)

Ptilinus fuscus (Anobiidae) was confirmed as the host species of Pelecotoma fennica (Rhipiphoridae, Pelecotominae). Females of Pelecotoma oviposit into the wood infested by the host larvae. First-instar larvae are elongate, unsclerotized, very different from the triungulinid larvae known in other rhipiphorids. They search actively for the host larvae in the wood; no phoresy is involved in the life cycle. The first-instar larvae temporarily paralyse the host larva and enter its body, overwintering (some perhaps more than once) as an endoparasite. In the spring of the year of emergence, the endoparasite engorges enormously (without moulting) and develops a special sclerotized caudal structure which is then used for perforation of the host's integument. The larva undergoes a further four ectoparasitic instars. The fifth (i.e. fourth ectoparasitic) instar differs considerably from the preceding three, and is capable of boring through the wood to prepare the emergence gallery for the adult. Pupation occurs in the wood. The rate of parasitization may locally far exceed 50%. Superparasitization by the first-instar larvae is possible, but usually only 1 larva survives to the ectoparasitic stage. Larval morphology of Pelecotoma is described and illustrated. Additional data on bionomics and larval morphology are also presented for the genus Metoecus (Rhipiphorinae). Comparing the biology and larval morphology of Pelecotoma with other Rhipiphoridae, it is assumed that ancestral rhipiphorids may have been xylophilous Tenebrionoidea with predaceous or omnivorous larvae, and that the ‘triungulinid’ larvae and phoretic habits may not belong to the groundplan of the family Rhipiphoridae. The widespread opinion considering Rhipiphoridae closely related to the family Mordellidae is questioned.     

Taxonomy and biology of final-instar larvae of some Eurytomidae (Hymenoptera: Chalcidoidea) associated with grasses in the UK

A comparative morphological study was made of the final-instar larval stage of 33 species of grass-inhabiting Eurytomidae belonging to four genera: Eurytoma Illiger, Tetramesa Walker, Sycophila Walker and Ahtola Claridge. Emphasis was placed upon chaetotaxy and mandibular structure. The final-instar larvae of the species are described, those of 26 species for the first time, and diagnostic characters and keys to genera and species are provided. Information, including new rearing records, is given on the larval biology of each eurytomid species, together with comments on the adult taxonomy of certain species.     

Partitioning phenotypic variation: genotypic,environmental and residual components from bryozoan skeletal morphology

Using morphometric studies of colonial (clonal) organisms such as Bryozoa grown as replicates in controlled laboratory experiments, phenotypic variation (hard part morphology) can be partitioned into its genotypic and environmental (ecophenotypic) components. The interaction between these, i.e. different genotypic responses to the same environmental change, can also be recognized. Palaeobiological studies are inherently constrained by species concepts based on morphotypes (preserved morphological phenotype). Uncertainties associated with fossil species concepts restrict the deductive resolution potential of fossil taxa in discussions of the broader biological questions of species evolution, ecology, biogeography and phylogeny. The relationship between species-level morphological variation and genetic variation in modern taxa is central to evaluating the viability of fossil morphotypes as biological species. Results from a preliminary study of three genotypes of Electra pilosa L., grown as replicate colonies in tanks comprising different microenvironmental conditions, allow for direct evaluation of morphospecies concepts. Numerical analyses (Cluster Analysis, Principal Component Analysis and Two-way ANOVA) of six morphometric characters demonstrated a strong Genotype control over zooid morphology and limited environmental (Tank) effects with minimal environmental differences among tanks. No significant Genotype by Tank (=environment) interactions were found for any characters. These results demonstrate that it is possible to quantify the extent to which a given trait is plastically expressed in different environments. The strong degree of correlation between morphology and genetics for this species is encouraging for the use of morphotypes as proxies for biological species.     

Identity of Mertensia oblongifolia (Nutt.) G. Don (Boraginaceae) and its allies in western North America

The current status of Mertensia oblongifolia (Nutt.) G. Don and its allied taxa is surveyed. On the bases of continuously coherent morphological characters and/or regionally correlated variations, more than 30 taxa, including species, subspecies, varieties, and 1 forma, previously considered different from M. oblongifolia , are now placed under synonymy of this species. Those taxa currently known as M. fusiformis Greene, M. bakeri Greene, and M. bakeri var. osterhoutii Williams are among the new synonyms. Typification, taxonomy, and morphological problems of M. oblongifolia are discussed.     

Intraspecific larval aggression in two species of Hyperini (Coleoptera: Curculionidae)

Two unusual types of behaviour (wandering and intraspecific aggressive behaviour) have been observed when rearing larvae of two species of the tribe Hyperini. Wandering of Hypera postica in search for food was observed in L1, L2 and L3 larvae, as well as in young L4 larvae. L1 larvae also disperse in response to crowding. Wandering of L2 to young L4 instars was a response to food shortage. Wandering of late L4 larva (‘prepupa’) occurs because of searching for a place to spin the cocoon and pupate. Encounters between the larvae may result in agonistic behaviour, and some larvae may die as a consequence of fighting. This aggression increases with food limitation. Agonistic intraspecific behaviour has not been described to date in weevil’s larvae and it probably may not occur under natural conditions when there is a plenty of food and larval densities are decreased by pathogens or parasitoids.     

Contrasting modes of handling moss for feeding and case‐building by the caddisfly Scelotrichia willcairnsi (Insecta: Trichoptera)

Final‐instar larvae of the microcaddisfly, Scelotrichia willcairnsi sp. nov. (Trichoptera: Hydroptilidae) were discovered feeding on a moss, Platyhypnidium muelleri in north‐eastern Queensland. Not only do the larvae feed on the moss, but their cases are constructed from fragments of moss leaves. Microscopic examination of larval guts and cases showed a total contrast in the angle at which the moss leaves are cut for each purpose: for cases the moss leaves are cut longitudinally, in parallel with the length of the cells, whereas for feeding the leaves are cut perpendicular to the leaf margin, across the cells, presumably an adaptation that releases the cell contents for digestion. The new caddisfly species is described based on the adult males. Scelotrichia willcairnsi represents the first Australian record of the South‐east Asian‐New Guinean Stactobiini genus Scelotrichia.     

Larval release rhythms of four species (family Grapsidae and Ocypodidae) of intertidal crabs on a coastal flat in western central Taiwan

Larval release rhythms of four crab species, Parasesarma tripectinis (Shen, 1940), Perisesarma bidens (De Haan, 1835), Hemigrapsus penicillatus (De Haan, 1835) and Baruna sinensis Tan and Huang, 1995, were studied in two different intertidal zones on a coastal flat. Of these, P. bidens is the only species that migrates between high and lower intertidal zones. In general, larval release patterns of these intertidal crabs are not too different from those of intertidal crabs living in other geographical regions, which often release larvae around new or full moons and nocturnal high tides. However, differences were apparent when compared with previous studies in that: (1) peak larval release periods of P. bidens occurred during submaximum tidal amplitudes, (2) a good portion of B. sinensis released their larvae during neap tides, (3) all four species released their larvae during both daytime and nighttime high tides although the percentage varied among these species, (4) three species released their larvae within 1 h after the time of high slack tide, but larval release of B. sinensis peaked between 1 and 2h after the time of maximum tidal height. Larval release pattern of Perisesarma bidens suggests that locomotory behaviour of high intertidal crabs may influence their larval release rhythms. Antipredatory adaptation may explain larval release rhythms of the four species examined in the present study. Larvae of Parasesarma tripectinis, P. bidens and Hemigrapsus penicillatus are more vulnerable to planktivorous fish predation and tend to release their larvae at nocturnal high tides during maximum tidal amplitudes. On the other hand, larvae of Baruna sinensis may be better adapted against planktivorous fish predation by having yellow-green colour. The timing of larval release of this species occurred throughout the lunar cycle and often at daytime 1-2 h after the time of high slack tide.     

Five invaginations and shedding of the larval epidermis during development of the hoplonemertean Pantinonemertes californiensis (Nemertea: Hoplonemertea)

We describe the planuliform larval development of the high intertidal hoplonemertean Pantinonemertes californiensis using confocal microscopy. We discovered that the newly hatched planuliform larva has two pairs of epidermal invaginations that disappear later in development. The anterior pair of invaginations has an apparent counterpart in other planuliform nemertean larvae; they give rise to the cerebral organs in at least two other hoplonemertean species. Although the adult P. californiensis possesses cerebral organs, we were not able to distinguish the anterior invaginations after 3 days of development. The developmental fate and homology of the posterior pair of invaginations is uncertain. We show that the proboscis develops in newly hatched larvae as a fifth distinct invagination just ventral of the apical plate. P. californiensis possesses a transitory larval epidermis, composed of approximately 80 multiciliated cells, which is normally shed at 3–4 days of development at 12–16°C.     

Ontogenetic and habitat-related changes in diet of late larval and juvenile suckers (Catostomidae) in Upper Klamath Lake, Oregon

We describe ontogenetic patterns in the diets of shortnose and Lost River suckers (15.8–92.8 mm standard length) from Upper Klamath Lake in summer 1999. Both species made a transition from surface and planktonic prey to benthic prey at about 20–30 mm standard length, corresponding to the approximate size of the juvenile morphological transition. Surface prey was dominated by adult chironomids and undigestable pollen, while benthic prey was dominated by larval chironomids, chydorids, and ostracods. In the 15–20-mm size class, pollen made up >75% of food particles in 68% of specimens, and only 2 specimens in this size class lacked any pollen grains. A better understanding of the prey selection process in larval suckers is needed to determine the importance of this potential source of starvation. The planktonic prey eaten during the surface-to-benthic feeding transition were widely distributed in the lake, suggesting that larvae use emergent vegetation primarily as a refuge from predators. On a numerical basis, crustaceans and larval chironomids contributed most to the diets of larger juveniles. In specimens >40 mm, shortnose suckers ate more Tanytarsini and Lost River suckers ate more chironomid pupae and chydorids. In specimens >40 mm there was a strong habitat-related difference in diet: specimens collected in offshore samples ate more chironomid larvae and pupae, harpacticoid copepods, and chydorids, whereas onshore specimens ate more cyclopoid copepods and invertebrate eggs. The transition from onshore to offshore habitat appears to occur over a broad size range of about 40–90 mm.     

The complete larval development of the crab Pilumnus spinifer (Brachyura: Xanthoidea: Pilumnidae) reared in the laboratory

The complete larval development of the crab Pilumnus spinifer from the western Mediterranean was obtained in the laboratory. All four zoeal stages and the megalopa are described and illustrated. The morphological characters of the larvae of Pilumnus spinifer are compared with those of other known larvae of the genus. The zoeae of P. spinifer show the rostral spine longer than the antennule (excluding aesthetascs); short lateral spines present on the carapace, and the mediolateral processes are present only in abdominal somites 2 and 3. The megalopa is similar to that of other Pilumnidae species. The morphology of the larval stages shows very similar characteristics to that of those of P. hirtellus and P. dasypodus, among the described stages of the family.     

Larval feeding habits of the Hepialidae (Lepidoptera)

The host-plant relationships of the Hepialidae are discussed in relation to host range and larval tunnelling behaviour. Larvae are mostly phytophagous on live angiosperms, gymnosperms, pteridophytes and mosses. Generally they specialize as feeders of leaf, stem/branch or root tissue, but dietary transitions from leaf to stem and root to stem occur in some species. An early period of mycophagy where larvae feed on fungi or dead decaying plant tissue is recorded in a wide range of species that feed on live embryophytes in later instars. Feeding patterns of the Hepialidae are compared with those of other major lepidopteran taxa and the evolutionary implications of hepialid feeding habits for the origin and derivation of larval host-plant relationships in Lepidoptera are discussed. It is suggested that the developmental transition from mycophagy to phytophagy in the Hepialidae is the result of partial ‘suppression’ of mycophagy in the generalist feeding ancestor of Lepidoptera.