Testing the directionality of evolution: the case of chydorid crustaceans

Although trends are of central interest to evolutionary biology, it is only recently that methodological advances have allowed rigorous statistical tests of putative trends in the evolution of discrete traits. Oligomerization is one such proposed trend that may have profoundly influenced evolutionary pathways in many types of animals, especially arthropods. It is a general hypothesis that repeated structures (such as appendage segments and spines) tend to evolve primarily through loss. Although largely untested, this principle of loss is commonly invoked in morphological studies of crustaceans for drawing conclusions about the systematic placements of taxa and about their phylogeny. We present a statistical evaluation of this hypothesis using a molecular phylogeny and character matrix for a family of crustaceans, the Chydoridae, analysed using maximum likelihood methods. We find that a unidirectional (loss-only) model of character evolution is a very poor fit to the data, but that there is evidence of a trend towards loss, with loss rates of structures being perhaps twice the rates of gain. Thus, our results caution against assuming loss a priori, in the absence of appropriate tests for the characters under consideration. However, oligomerization, considered as a tendency but not a rule, may indeed have had ramifications for the types of functional and ecological shifts that have been more common during evolutionary diversification.     

Preface: The phylogeny, fossil record and ecological diversity of ostracod crustaceans

After more than two centuries of research, more than 65,000 living and fossil ostracod species have been described and studied, yet much remains to be learned about this ancient, widespread and diverse group of bivalved arthropods. Their higher classification and phylogeny are subjects of vigorous debate, as is their position in the broader picture of crustacean phylogeny. At the same time, major advances in our understanding of ostracod lineages and their relationships are resulting from the application of innovative approaches and techniques. This preface provides a contextual overview of the 15 contributions to this volume, which resulted from the 14th International Symposium on Ostracoda (ISO2001) held in 2001at Shizuoka, Japan. As such it provides a cross-section of topics at the forefront of research on the evolution and diversity of Ostracoda, and indicates directions for future work.     

Anaerobic capacity of a crustacean sensitive to low environmental oxygen tensions,the freshwater amphipod Gammarus pulex (L.)

We investigated the accumulation of haemolymph L-lactate in response to anoxia, progressive hypoxia and activity, in a species that rarely encounters low PO₂ in its natural environment, the freshwater amphipod Gammarus pulex (L.). Individuals survived <1.5 h anoxia and accumulated L-lactate in the haemolymph at a rate of 10.26 mmol l^–1 h^–1. No lactate was excreted into the medium during this time. Despite the fact that most individuals of G. pulex were oxyconformers, exposure to acutely declining PO₂s did not result in a switch to anaerobic pathways until PO₂ < 4.4 kPa. Even then the concentrations accumulated remained very low until PO₂ < 0.46 kPa. There was no accumulation of L-lactate in the haemolymph as a result of exhaustive activity. Consequently, it was suggested that (a) it cannot be automatically assumed that anaerobic metabolism is switched on only at the point at which oxyregulation breaks down, (b) that the poor anaerobic capacity observed plays little, if any, role in the metabolic response to hypoxia by Gammarus pulex.     

On the influence of fishes on the evolution of benthic crustaceans

Though the interweavement of trophic chains in marine ecosystems of the past cannot be reconstructed, as the abundance, nutrition, reproductive rates and other aspects of the biology of fossil species are not known, it is possible to correlate trends in the evolution of single groups of animals, in the presented case the Crustacea, with the influence of other evolving organisms. The evolution of carnivorous, buoyant, well swimming modern fishes probably induced profound changes in marine ecosystems and influenced the evolution of crustaceans. Indications for these interactions are a. the absence or rare occurrence of defenseless archaic crustaceans in habitats, which are populated by teleostean fishes and the survival of some of these forms in refuges like caves and subterranean waters, and b. the reduction of the pleon in the course of the mesozoic evolution of the Decapoda, which occurs parallel to the radiation of the Teleostei, namely in the period between the early Jurassic and the Tertiary. The shortening of the pleon is interreted as a consequence of the stepwise change from a hyperbenthic to a more benthic Iife-stJe and the abandonment of the caridoid escape reaction. Other adaptations are also construed as results from the selective pressure produced by predators. Extant crustaceans which are able to coexist with fishes, among the macrozoobenthos especially the Decapoda and Peracarida, have a variety of protective adaptations, which help to reduce predation.     

Diapause of copepods as an element for stabilizing the parasite system of some fish helminths

The analysis of infection dynamics of copepods, which are the intermediate hosts of three helminth species of freshwater fishes (Triaenophorus crassus, Proteocephalus exiguus and P. percae), has shown that diapause in the life cycle of the copepods is favourable for preserving the infection in the waterbody until physiological prerequisites for successful infection of the final host are acquired. The peculiarity of a copepod's life cycle may determine the strategy of a parasite in its preimaginal phase as a waiting stage, and the duration of the residence of helminth larvae in copepods that have an obligatory diapause, is one of the elements the provide stability in parasite systems.     

Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater

The isopod crustaceans are diverse both morphologically and in described species numbers. Nearly 950 described species (∼9% of all isopods) live in continental waters, and possibly 1,400 species remain undescribed. The high frequency of cryptic species suggests that these figures are underestimates. Several major freshwater taxa have ancient biogeographic patterns dating from the division of the continents into Laurasia (Asellidae, Stenasellidae) and Gondwana (Phreatoicidea, Protojaniridae and Heterias). The suborder Asellota has the most described freshwater species, mostly in the families Asellidae and Stenasellidae. The suborder Phreatoicidea has the largest number of endemic genera. Other primary freshwater taxa have small numbers of described species, although more species are being discovered, especially in the southern hemisphere. The Oniscidea, although primarily terrestrial, has a small number of freshwater species. A diverse group of more derived isopods, the ‘Flabellifera’ sensu lato has regionally important species richness, such as in the Amazon River. These taxa are transitional between marine and freshwater realms and represent multiple colonisations of continental habitats. Most species of freshwater isopods species and many genera are narrow range endemics. This endemism ensures that human demand for fresh water will place these isopods at an increasing risk of extinction, as has already happened in a few documented cases. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Monoaminergic neurons in the nervous system of crustaceans

Summary Certain neurons in the nervous system of the malacostracan crustaceans give rise to a predominantly green and a sparse yellow fluorophore in the histochemical fluorescence method of Falck-Hillarp. The same applies to the whole of Crustacea. The green fluorophore is probably a catecholamine; the yellow to brown-yellow has not yet been identified.The biogenic amine responsible for the green fluorescence, besides being found in diffusely distributed fibres, also appears in distinct areas of fibre concentrations in the central nervous system. The protocerebrum of the malacostracans contains three areas: the central body and two areas in the top of the brain, one anterior and one posterior. The latter two are not recognized as separate areas in ordinary histological preparations. In addition, the optic neuropiles are fluorescent, some with a distinct stratification of the fluorophore. The deuto and tritocerebrum and the ventral nerve cord also contain monoaminergic neurons. Of the brightly fluorescent areas in the whole of Crustacea, only the central body consistently exists in all species. The other areas of concentrated fluorescent neuropile are restricted to smaller taxonomic units and differ from each other. p The monoaminergic neurons in Crustacea are sensory, motor, and internuncial, and also belong to a fourth type which mimics the neurosecretory neurons in neurohaemal organs. Only one example of a monoaminergic sensory neuron is known (in Anemia, a non-malacostracan, Aramant and Elofsson 1976), a few motor and a few neurosecretory mimics (the latter in malacostracans). Most are internuncials. Acknowledgement. We have enjoyed the laboratory facilities at the Department of Histology, Faculty of Medicine, and express our sincere thanks to Prof. Bengt Falck.-Grants from the Swedish Natural Science Research Council (2760-007) and the Swedish Medical Research Council (04X-712) supported the work     

Diapause development in the chestnut weevilCurculio elephas

Larval diapause development in the chestnut weevilCurculio elephas (Coleoptera, Curculionidae) was studied in the laboratory at different temperatures. The results proved that exposure to low temperatures (3–6°C) in the period December–February is not required to complete diapause. The diapause is terminated in December and from January on, the larvae can initiate post-diapause morphogenesis in the laboratory, if temperatures allow it. In the field developmental rates are negligible during winter cold (4–6°C) and only after March morphogenesis can proceed with no interruption until adult emergence. Diapause and post-diapause quiescence contribute to individual synchronization for initiation of development. The observed spread of adult emergence was 30 days in the laboratory. This variability produced during post-diapause development may be a response to annual variation in the phenology of the chestnuts.     

Genetic diversity in populations of a freshwater ciliate

RAPD fingerprinting with nine different primers revealed that all of 18 E. aediculatus isolates from nine ponds and streams in western Germany, France and the U.S.A. were genetically different. The extent of genetic similarity between genotypes from different waters did not show a significant relationship with the geographical distance among habitats, although genotypes isolated from the same habitat showed a higher genetic similarity than genotypes isolated from different habitats. Phylogenetic analyses of RAPD patterns indicate a separation of E. aediculatus strains into subgroups within one species, but all strains were genetically more similar to one another than to strains from two other Euplotes species. Crossings of the different E. aediculatus strains revealed they belonged to seven mating types of one gene pool. The high genetic diversity observed is explained by a frequent occurrence of conjugation in the studied populations.     

Limb morphogenesis in the branchiopod crustacean, Thamnocephalus platyurus, and the evolution of proximal limb lobes within Anostraca

Crustacean limbs exhibit highly diverse morphologies. One major route of diversification is in the number and position of branches arising from the proximal part of the limb. Here I describe development of larvae of the branchiopod crustacean, Thamnocephalus platyurus and describe in detail the development of the thoracic limbs. The thoracic limbs bear proximal branches both medially and laterally. The most proximal branches on either side (gnathobase and pre-epipod) show a similar developmental history: they develop via fusion of two rudiments into a single adult branch. However, phylogenetic analysis suggests that the developmental fusions have distinct evolutionary histories. In one case (gnathobase), the developmental rudiments reflect the ancestral adult morphology of two distinct branches. In the other (pre-epipod), the rudiments are an apparent novelty within the Anostraca and develop into two adult structures in only a single derived family.     

Diapause, quiescence, hatching requirements: what we can learn from large freshwater branchiopods (Crustacea: Branchiopoda: Anostraca, Notostraca, Conchostraca)

The extent to which dormancy in large freshwater branchiopods is controlled endogenously (diapause) or exogenously (quiescence) is not always clear. It is assumed that both processes occur even within the same brood. Based on the effectiveness of common diapause-deactivating processes such as desiccation, hibernation, and resting, it can be stated that diapause is not a general process controlling responsiveness of large freshwater branchiopod eggs. Only in limited cases unequivocal evidence for the positive influence of these treatments is found.With few exceptions, hatching of activated cysts is effected by specific conditions that may even differ among conspecific populations. Generally, each species (or even population) has a specific temperature range or regime for optimal hatching performance. In a suitable thermal environment with sufficient light and oxygen, hatching is invariably invoked by a low osmotic medium.The erratic hatching pattern in most species is thought to be an adaptation to the variable temporary habitat. Hatching is generally spread over several days or even weeks, but the highest peak usually occurs on the first or second day of hatching. Low hatching percentages were found only in subtropical/desert species and may be a reflection of the low chances for successful reproduction. Generation carry-over of propagules in the egg bank by dormancy, and hatching at low conductivity, are, together with obligate oviparity and absence of an asexual life cycle phase, thought to be highly adaptive to the temporary environment. This enabled large branchiopods to survive since the Upper Cambrian.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

The molecular diversity of the methanogenic community in a hypereutrophic freshwater lake determined by PCR-RFLP

AIMS: To combine database-held sequence information with a programme of experimental molecular ecology to define the methanogenic community of a hypereutrophic lake by a PCR-restriction fragment length polymorphism (RFLP) analysis. METHODS AND RESULTS: Methanogen diversity in a hypereutrophic freshwater lake was analysed using 16S rDNA PCR-RFLP. Database-held 16S rRNA gene sequences for 76 diverse methanogens were analysed for specific restriction sites that permitted unequivocal differentiation of methanogens. Restriction digestion and agarose gel electrophoresis of the 16S rDNA from selected methanogen pure cultures generated observed restriction profiles that corroborated the expected patterns. This method was then tested by analysing methanogen diversity in samples obtained over 1 year from sediment and water samples taken from the same sampling site. CONCLUSIONS: Restriction analysis of the 16S rRNA gene sequences from 157 methanogen clones generated from lakewater and sediment samples showed that over 50% were similar to Methanoculleus spp. Furthermore, a total of 16 RFLP types (1-16) were identified, eight of which contained no cultured representative archaeal 16S rRNA gene sequences. SIGNIFICANCE AND IMPACT OF THE STUDY: This RFLP strategy provides a robust and reliable means to rapidly identify methanogens in the environment.     

Monoamine-containing neurons in the optic ganglia of crustaceans and insects

Summary With the fluorescence method of Falck and Hillarp, the presence and localization of monoaminergic neurons in the optic ganglia of several crustaceans and insects have been investigated. It was found that in both classes the monoaminergic terminals, when present, appeared (especially in the medullae externa and interna of the crustaceans and the medulla of the insects) in strata specific for each species. So far, the only monoamine (visualized by this technique) present in the crustacean optic ganglia is dopamine, whereas in the Insecta, the catecholamines dopamine and noradrenaline, and the indolamine, 5-hydroxytryptamine, are found in the optic lobe. But in the Insecta, different species show different content of these amines.This work was supported by grants 2760-3 and 2760-4 from the Swedish Natural Science Research Council (R.E.), by a fellowship from Deutsche Forschungsgemeinschaft, and a grant from the Swedish Medical Research Council B72-14X-712-D7B (N.K.). We are very grateful to the director of the Department of Histology, Faculty of Medicine, Lund, Professor Bengt Falck, who put all his facilities and knowledge at our disposal.     

Species diversity of freshwater hyphomycetes in some streams of Pakistan. Comparison of sampling techniques

Freshwater hyphomycete communities in four streams were sampled by filtration of water samples, trapping conidia in artificial foam, and examination of alder leaf pack baits and random sampling of naturally occurring submerged leaves. No two communities of freshwater hyphomycetes detected by different sampling techniques used singly in a stream showed 100% similarity. Based on relative frequency values, the same 10 top-ranking species in a stream were detected by artificial foam trap and on the baited leaves. These species differed in ranking, and the community in artificial foam was riche than on baited leaves. Of the various communities detected by different sampling techniques used singly or in combination, those detected by artificial foam trap or a combination of this technique with leaf baiting or with leaf baiting and randomly sampled leaves or filtration showed the highest similarity to the total community based on data generated using four sampling techniques simultaneously.This work was conducted under Grant No. PSF Project Environment 3 from the Pakistan Science Foundation.     

Diapause in a large aggregation of a tropical beetle

Abstract. 1_˜ Diapause is described for adults of Stenotarsus rotundus Arrow (Endomychidae, Coleoptera) in the tropical lowland forest on Barro Colorado Island, Panama.
2 . Diapause lasted up to 10 months each year, covering some 6 months of the wet season plus 4 months of the dry season. When in diapause the beetles clustered in an aggregation of up to 70 ,000 individuals on the stem of a palm tree, the same individual palm tree in at least four successive years.
3. The habitat and habits of the beetle when not in diapause are unknown, but one or two generations may be produced during this period. Beetles returned to the diapause site from June until late August.
4. Soon after arrival at the diapause site the flight muscles degenerated and did not develop again until February/March. The ovaries were undeveloped when the beetles arrived at the site and remained in that condition until the onset of yolk deposition in March. At the beginning of the rainy season in April the beetles mated and dispersed.
5. Lean dry weight of the beetles did not decrease noticeably during diapause, but fat content did. It is suggested that the beetles do not feed, or feed very little, while in diapause. During diapause, the metabolic rate is surprisingly low considering a mean ambient temperature of 25° C.
6. Juvenile hormone analogue (ZR515) stimulated flight muscle and ovary development. Daylength, when different from LD 12: 12 , initiated a partial flight muscle development, but there was no effect on the ovaries. Field observations suggest that neither temperature nor humidity had an effect on the termination of diapause.
7. Although the development of both flight muscles and ovaries were stimulated by juvenile hormone analogue and both developed around March of each year, it is suggested that the mechanisms causing this development are different.     

Catecholaminergic innervation of muscles in the hindgut of crustaceans

Summary The crustacean species Pacifastacus leniusculus and Gammarus pulex were investigated by electron microscopy in a search for possible neuromuscular junctions in the hindgut, which has a rich supply of catecholaminergic fibres. True neuromuscular synapses were found in both species between nerve terminals containing dense-core vesicles (80–110 nm in diam.) and muscle fibres. We suggest that the dense-core vesicle terminals contain a catecholamine, and this is supported by ultrahistochemical tests for monoamines. Two types of junctions are found: one in which the nerve terminal is embedded in the muscle cell (both species) and one in which protrusions from the muscle cell meet nerve terminals (Pacifastacus). Gammarus pulex, which has only circular muscles in the hindgut, has only catecholaminergic innervation, whereas Pacifastacus leniusculus has circular and longitudinal muscles both with at least two types of innervation.The investigation was supported by grants from the Swedish Natural Science Research Council (B 2760-009), the Hungarian Academy of Sciences, the Royal Swedish Academy of Sciences, and the Magnus Bergvall Foundation. We are also indebted to Mrs. Lena Sandell for her skilful technical assistance     

On Indiacypris luxata (Brady), a freshwater ostracod (Crustacea: Entomostraca) from Sri Lanka

Indiacypris luxata (Brady), a freshwater ostracod from Sri Lanka is described and both the valves and the limbs and soft parts are figured.     

Species diversity of freshwater hyphomycetes in some streams of Pakistan. III. Autumnal colonization of freshwater hyphomycetes on bait leaves

Significantly more species of freshwater hyphomycetes colonized bait leaves of alder (16 species) and willow (14 species) during early submersion period (1 wk) than those of oak (8 species). Production of conidia was also higher on alder and willow leaves than on oak leaves. Higher amounts of dry mass were lost from the alder (28.6%) and willow leaves (29.4%) than from the oak leaves (18.7%). Rapid loss of dry mass was accompanied by rapid development of a fungal community. Fungal species took a longer time (6 wk) to reach their peak of occurrence on oak than on willow and alder leaves (3–4 wk).Flagellospora curvula dominated the assemblages of freshwater hyphomycetes on alder and willow leaves, andLunulospora curvula dominated the assemblage on oak leaves. The freshwater hyphomycete community showed higher values of species diversity (H value) on oak leaves than on alder and willow leaves.