Three species of Isotoma (Collembola, Isotomidae) based on morphology, isozymes and ecology

Morphological markers and isozymes were used for identifying three presumed species of the Isotoma genus. Morphological traits separated three taxa of the genus. 10 isozymes determined by at least 11 loci were analysed from each taxon, and 2 loci were taxon-specific, supporting the hypothesis that the three taxa represented three species. The genetic variation found within the taxa measured as fraction of polymorphic loci at the 99% level was 0.82, 0.55 and 0.55 with the corresponding observed heterozygosity 0.15, 0.09 and 0.12. Two populations of the same taxon from Denmark and Norway, respectively, were very closely related. Additional ecological criteria, obtained from a literature survey, also revealed pronounced differences between the three taxa. Due to these facts three distinct species are proposed, namely I. anglicana Lubbock 1862, I. riparia Nicolet 1841 and I. viridis Bourlet 1839.     

A novel method of self-righting in the springtail Sminthurus viridis (Insecta: Collembola)

Photographic evidence shows that the ventral tube can be used as a tentacle-like mechanical aid for self-righting from an inverted position. It is proposed that the extraordinary length of the vesicles of the ventral tube in globular springtails has been evolved in order to combat postural instabilities resulting from spherical symmetry of body form.     

Temperature-controlled under-water egg dormancy and postflood hatching in Isotoma viridis (Collembola) as forms of adaptation to annual long-term flooding

Summary Incubation experiments with eggs of a population of Isotoma viridis, which is exposed to annual long-term flooding from about April to July, as well as field observations show that temperature controls both, underwater egg dormancy and immediate postflood hatching. The population is located at the Eder Freshwater Reservoir in Germany.If constant experimental temperatures are above 14°C, almost all eggs are nondormant. Dormancy is established at temperatures below 15°C, but embryonic development is completed. Experiments indicate that of the environmental factors that change drastically at the end of submergence (light, turgor pressure, oxygen, a.o.), only temperature acts as a hatching trigger. Hatching of the previously dormant eggs occurs at a constant threshold temperature of 16°C, mainly within 2 to 20 days after temperature elevation, but most of these eggs need even higher temperatures to hatch. Remaining eggs were partly stimulated to hatch by recooling them at 7°C for some days and then rewarming them again.The threshold temperatures observed are unusually high for Collembola and seem to be the result of selection by the special floodplain conditions. During normal years, the surface temperatures of submerged soil usually do not exceed threshold limits before summer drainage. This allows both, protection from under-water hatching and an optimal timing of hatching at the very beginning of the main terrestrial period. The experiments show that above the threshold temperature (in warm summers), individuals can hatch under water and survive submerged for 10–15 days. They can survive even longer in the water habitat, if emergent structures enable them to climb onto the water surface. Furthermore, a considerable polymorphism observed in some hatching properties improves the chance to survive under the unpredictable floodplain conditions.     

Two new Megalothorax species of the minimus group (Collembola,Neelidae)

Two new Megalothorax species, Megalothorax potapovi sp. n. from the Russian Far East and Megalothorax sanguineus sp. n. from the French Pyrénées are described. The two new species have a set of morphological characters (including a smooth mucro) that places them among the minimus group sensu Schneider and D’Haese (2013). Megalothorax potapovi characteristics include dorsal protuberance on forehead, peculiar chaetotaxy of antenna III and strong lanceolate chaetae on body. Megalothorax sanguineus characteristics include strong red pigmentation, large network of integumentary channels on head and elongated apex of the two postero-distal spines of dens. The DNA barcodes ( cytochrome oxidase subunit I–COI) of the two species are also provided and analyzed among a broader sampling of the genus in order to support further their specific status. A special focus is given to the labral morphological characteristics. Pseudopores-like elements are reported for the first time in the genus. Positions of the τ-chaetae near the dorsal sensory field of thorax II are compared for several species of the genus.     

Das Doppelauge vonEntomobrya muscorum Nicolet (Insecta,Collembola)

Zusammenfassung Der bodenlebende CollembolaEntomobrya muscorum hat an jeder Kopfseite ein loses Aggregat von acht Ommatidien, von denen zwei einen deutlich kleineren Linsendurchmesser aufweisen. Die sechs großen Ommatidien werden als Haupt-, die zwei kleinen als Nebenaugen bezeichnet. Beide Ommatidientypen haben verschieden gebaute Rhabdome, so daß der Komplex ein Doppelauge darstellt. Es wird der Bau der Hauptaugen geschildert. Ihr sensibler Teil setzt sich aus acht Retinulazellen zusammen, die ein offenes Rhabdom in zwei Lagen bilden. Eine zentrale sechste Zelle in der distalen Schicht weicht in ihrer Feinstruktur deutlich von den übrigen ab. Die Rhabdome beider Ommatidientypen werden miteinander verglichen, und ihre funktionelle Bedeutung vor allem in Hinblick auf eine mögliche Arbeitsteilung für eine Polarisationswahrnehmung wird erörtert. Es wird kurz die phylogenetische Bedeutung dieser Augen für die Insekten und die Collembolen selbst angeschnitten.

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The double eye ofEntomobrya muscorum nicolet (Insecta, Collembola)

Summary The ground living CollembolaEntomobrya muscorum has on each side of the head eight circular ommatidia in an arrangement characteristic of the species. Of these two have a distinctly smaller lens. The six big ommatidia are called primary eyes, the two small ones accessory eyes. The two types of ommatidia have different rhabdom structures, so justifying the term double eye. The structure of the primary eyes is described. The photosensitive part is constructed of eight retinula cells which form an open rhabdom in two layers. The central sixth cell in the distal layer is quite different from the others. The rhabdoms of both types are compared. The possible function of the accessory eyes in perception of the e-vector is discussed. The phylogenetic significance of these eyes for the Insecta and the Collembola is considered.

     

Molecular evidence for cryptic species within Cylicostephanus minutus (Nematoda: Strongylidae)

The nucleotide sequences of the first and second internal transcribed spacers of nuclear ribosomal DNA were determined for adults of Cylicostephanus minutus from different geographical origins. The lengths of first and second internal transcribed spacer sequences ranged from 370 to 372bp and 215 to 216bp, respectively. Pairwise sequence comparisons revealed that some individuals of C. minutus had identical first and second internal transcribed spacer sequences, whereas others differed by 3.0% and 7.4% in their first and second transcribed spacers, respectively. Some individuals with sequence differences originated from the same host. The levels of difference within C. minutus were higher than that between the morphologically distinct species, Cylicostephanus goldi and Cylicostephanus longibursatus (0.8% for the first internal transcribed spacer and 3.8% for the second internal transcribed spacer). The data provide support for the proposal that C. minutus represents a complex of at least two species. In order to study the population genetic structure of C. minutus, a PCR-linked single-strand conformation polymorphism technique was also established.     

Ultrastructure of the male accessory glands ofAllacma fusca(Insecta, Collembola)

The accessory glands ofAllacma fusca(L.) (Insecta, Collembola, Sminthuridae) consist of a series of secretory units that are arranged in parallel and open into the ejaculatory duct. Each unit is composed of microvillate cells stacked around a common cavity. Basal cells are involved in ion-control of fluids from the hemocoel to the cavity. The intermediate and apical cells, which have a laminar appearance and contain many microtubules, are involved in the structural integrity of the unit. Supporting cells ensheath the most apical cells. Large openings in the cuticle allow the gland secretion to flow into the ejaculatory duct lumen. These openings are protected by a porous cuticle different from that lining the epithelium of the ejaculatory duct. Conspicuous muscle fibers run along the lateroventral side of the ejaculatory duct beneath the insertion of the accessory glands. The fine structure of the accessory glands indicates that they are type I ectodermic glands as defined by Noirot & Quennedey (1974). Their function could be to control the fluidity of the material for spermatophore formation and to ensure the proper physiological conditions for spermatozoa stored in the ejaculatory duct lumen.     

Molecular evidence for further overlooked species within the Gammarus fossarum complex (Crustacea: Amphipoda)

The freshwater amphipod Gammarus fossarum Koch, in Panzer, 1836 is a locally abundant keystone species mainly occuring in European headwaters but also in larger rivers. Genetic studies in the past 25 years have revealed three cryptic species within nominal G. fossarum (types A, B and C). Assignments of specimens to these types were based on allozyme and 16S markers. Today, a fragment of the cytochrome c oxidase subunit 1 (CO1) is primarily used as a genetic marker for species assignments (‘DNA Barcoding’), yet not a single CO1 sequence of G. fossarum is available in the Barcode of Life Database. We analysed new CO1 and 16S data for German, Hungarian and Croatian G. fossarum specimens and compared them with 16S and CO1 sequences of G. fossarum from GenBank. Thereby, we close the gap between traditional allozyme- and 16S-based species assignments and modern CO1 barcoding. Studying genetic variation in 55 specimens from 29 populations, we identified between 11 and 23 novel and genetically distinct clades using distance- and tree-based methods. Our results suggest that G. fossarum comprises several additional, yet unrecognised, species in particular from the Balkan region. Therefore, a taxonomic revision and biogeographic reconsideration of the G. fossarum complex is urgently needed.     

Molecular evidence of cryptic species within the Liriomyza huidobrensis (Diptera: Agromyzidae)

Phylogenetic relationships among populations of the polyphagous pea leafminer, Liriomyza huidobrensis (Blanchard), were investigated using DNA sequence data. Maximum parsimony analysis of 941 bp of mitochondrial cytochrome oxidase I and II genes showed that L. huidobrensis contains two well-defined monophyletic groups, one composed of specimens from California and Hawaii and one composed of specimens from South and Central America together with populations that have been recently introduced into other parts of the world. The differentiation between the two clades within L. huidobrensis is equivalent to that seen between other agromyzid species, suggesting that L. huidobrensis as currently defined contains two cryptic species. This finding is consistent with field observations of differences in pest status and insecticide resistance between L. huidobrensis populations. Until additional studies are complete, no changes in L. huidobrensis taxonomy are proposed. However, researchers and quarantine officials may wish to consider the findings of the current study in designing research, pest management, and quarantine programs for L. huidobrensis.     

Le comportement reproducteur de Sminthurus viridis (L.) (Collembola,Symphypleona)

Different phases of the reproductive behaviour of Sminthurus viridis are analysed. Stalked spermatophores are deposited by the ♂, but the sperm is transmitted during a primitive courtship. Oviposition consists of characteristic sequences, present in all species which deposit coated eggs. Here, however, the shortness of the sequences results from the eggs being laid in batches and from the frequently dry conditions in the natural biotope of this species. The anatomy of the genital organs of ♂♂ and ♀♀ is described, with a particular study of the genital plate of the ♂ by SEM.     

Molecular and morphological evidence for multiple species within Paranoplocephala omphalodes (Cestoda, Anoplocephalidae) in Microtus voles (Arvicolinae)

Haukisalmi, V., Wickström, L. M., Henttonen, H., Hantula, J. & Gubányi, A. (2004). Molecular and morphological evidence for multiple species within Paranoplocephala omphalodes (Cestoda, Anoplocephalidae) in Microtus voles (Arvicolinae). —Zoologica Scripta, 33, 277–290. The present study was designed to test the hypothesis that the anoplocephalid cestode Paranoplocephala omphalodes (Hermann, 1783), a Holarctic parasite of Microtus voles, is a complex of host‐specific species, rather than a single host‐generalist species, using uni‐ and multivariate morphometrics and DNA sequence data from the mitochondrial cytochrome oxidase I gene. The phylogenetic methods applied to the mtDNA sequence data showed consistently that the cestodes morphologically recognizable as P. omphalodes include four well‐supported monophyletic groups, representing at least three distinct, largely host‐specific species. Multivariate morphometrics (discriminant analysis) successfully distinguished the four main mtDNA clades of P. omphalodes‐like cestodes. The true P. omphalodes is shown to be a parasite of Microtus arvalis, M. agrestis and Clethrionomys glareolus in Europe. Microtus oeconomus harbours two host‐specific, allopatric and possibly conspecific clades, one with a Holarctic and another with an (eastern) Beringian (Alaskan) distribution. The eastern Beringian endemic M. miurus is also parasitized with a host‐specific, morphologically divergent species of Paranoplocephala. The cestode clades recognized in M. oeconomus and M. miurus represent 2–3 undescribed species. Molecular phylogenetic analyses supported the monophyly of the ‘northern clade’ of Paranoplocephala spp., an assemblage including P. kalelai from Clethrionomys spp., P. macrocephala from Microtus spp. and all clades of P. omphalodes‐like cestodes except those representing the true P. omphalodes from Europe. The intra‐ and interspecific phylogeny within the northern clade is compared tentatively with the known evolutionary history of the hosts.     

Dicyrtotnina and Ptenothrix (Insecta:Collembola) of the Solomon Islands

This paper deals primarily with the collembolan species of the subfamily Dicyrtominae collected during the Royal Society Expedition to the Solomon Islands of 1965 by P. N. Lawrence, supplemented by material from these islands collected by Dr and Mrs P. J. M. Greenslade, all of which is in the British Museum (Natural History). Additional supporting material described here has been derived from the Noona Dan Expedition of the Zoological Museum, Copenhagen and from the Bernice P. Bishop Museum, Hawaii. I should like to express my thanks to the individuals and Museum sconcerned.     

Endosymbiotic microorganisms in Adelges (Sacchiphantes) viridis (Insecta,Hemiptera, Adelgoidea: Adelgidae): Molecular characterization,ultrastructure and transovarial transmission

The aim of this paper was to identify endosymbiotic microorganisms living in the body cavity of a Polish population of an aphid, Adelges (Sacchiphantes) viridis, as well as to describe their ultrastructure and mode of transmission between generations. Molecular data (amplification and sequencing of 16S rRNA genes) indicated that endosymbionts of A. (S.) viridis are Betaproteobacteria of the species “Candidatus Vallotia virida”. Endosymbiotic bacteria are rod-shaped and localized in the cytoplasm of specific cells, termed bacteriocytes, of host insects. Endosymbionts sharing the same bacteriocytes differ in the density of their cytoplasm. There are two morphotypes of endosymbiotic bacteria: with electron-dense cytoplasm and electron-translucent cytoplasm. Since only bacteria containing electron-dense cytoplasm were observed in the binary fusion stage, differences in density of the cytoplasm are probably due to changes in the cytoskeleton of bacteria during division. Endosymbionts of A. (S.) viridis are transovarially (i.e. via oocytes) transmitted from the mother to the offspring.     

A New Species of Halophilous Collembola (Insecta) from Korea

Three species of interstitial Collembola in Entomobryidae were identified from South Korean coast. One of them, Sinella (Coecobrya) maritima, is new to science. So far 21 halophilous species in 10 families of Collembola, including the new species in this paper, were reported from the Korean peninsula.     

Studies on populations of Folsomia candida (Insecta: Collembola)

Summary Experiments are described that were designed to investigate the effects of food availability and rate of exploitation on the growth and production of populations of Folsomia candida (Willem). In an initial experiment in which there was excess food it was found that increasing the rate of exploitation resulted in increases in both the numerical and biomass productivity of the populations. In a second experiment it was shown that, when there is severe competition for food, the rate of exploitation does not affect either the biomass or the numerical production. It is concluded that the effect of overcrowding, in the form of competition for space, does contribute to the growth in numbers of populations, but that the supply of food plays a more important role in regulating the population. Anomalous results, showing that exploitation has a lesser effect when there is severe competition for food, are discussed.     

High diversity and community permeability: the riparian Collembola (Insecta) of a Pyrenean massif

This work deals with the structure of Collembolan communities in riparian habitats. Sixty samples collected along running waters in a Pyrenean massif have been analysed. Diversity was much higher than for any comparable habitat studied in the literature. The proportion of rare species was particularly high, and explained a large part of the observed diversity. The contribution of specialized hydrophilous species to overall diversity was low: they were 5 times less numerous than non-hydrophilous species for a similar global abundance. Correspondence analysis showed that the riparian community was loosely structured, without any strong determining factor. Epigeomorphic species of hydrophilous Collembola were, however, clustered in a well-defined group. The forest type, of major importance for soil fauna diversity in the area, was a poor predictor of diversity in the riparian habitats. Conversely, richness was significantly related to distance from water, in spite of the samples having been all collected from permanently water-saturated substrates. The importance of non-hydrophilous species for the diversity of riparian habitats along running water is interpreted as reflecting the ecological permeability of the hydrophilous community, resulting from the spatial organisation of the habitat in narrow strips, and its frequent disturbance by flooding. Finally, the riparian habitat may act as a refuge for a significant proportion of the soil species affected by reafforestation, presently the most severe disturbance of Pyrenean ecosystems.     

Assessing species boundaries and evolutionary relationships in a group of south-western European species of Isotomurus (Collembola, Isotomidae) using allozyme data

In this study allozyme data were used to examine evolutionary relationships in the cosmopolitan collembolan genus Isotomurus . Fifty-six south-western European populations from 13 species were compared, with the aim of delimiting species boundaries and testing the robustness of the most frequently used morphological characters for species identification. Observed allozyme variability suggests that genetic differentiation is more pronounced than the amount of morphological variability among and within species. Convergence in models of pigmentation pattern may lead to erroneous taxonomic identification when this is the only feature used for species diagnosis. In this respect, this study confirms that diagnosis is greatly improved when assisted by biochemical or molecular analyses. Allozyme data have also been used to reconstruct evolutionary hypotheses for relationships at the species and population level. The monophyly of Isotomurus maculatus , I. unifasciatus , I. fucicolus , I. nebulosus and I. pseudopalustris was confirmed on molecular grounds. Conversely, the monophyly of I. ghibellinus and I. palustris was rejected, suggesting the presence of cryptic species.     

Polymorphism of polytene chromosomes in Bilobella aurantiaca (Insecta: Collembola)

Polytene chromosomes in the salivary glands of Bilobella aurantiaca are analysed in a population from Sierra de Gredos (Central Spain). They display a high degree of heterozygosity. Only a short part of the total length of the seven pairs of homologous chromosomes, examined in a single favourable specimen, was homozygous. Concerning the short arm of chromosomes III and IV, the respective proportion of heterozygosity for the whole population was 38.6% and 92.9%. Some hypotheses are formulated to account for these results: differential funtioning of homologous chromosomes, generalisation of minute duplications and deficiencies, complex inversions (which cannot be specified owing to the asynapsis of homologous chromosomes), and possibly local large deletions.