Blastodermic cuticles of a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The formation and structure of the blastodermic cuticles of a springtail, Tomocerus ishibashii Yosii (Collembola : Tomoceridae) are described together with the change of egg membrane. The blastodermic cuticles of the Collembola are 2-layered, and formed in the early stages of the embryonic development, preceding the differentiation of germ band. The first blastodermic cuticle is thicker (about 0.8-1.5 μm in thickness) and its surface is provided with complex structures, whereas the second one is thinner (about 0.2-0.4 μm in thickness) and smooth. About 3 days after oviposition, the chorion (about 2 μm in thickness) splits into 2 and the first blastodermic cuticle, provided with many projections and 4 large spines appear on the surface of the egg. Three types of projections are distinguished: button-, cone- and seta-like structures. The halves of the ruptured chorion are attached to the first blastodermic cuticle on both sides below the spines, and no projections are found in the regions concealed by the ruptured chorion. The projections of the first blastodermic cuticle are formed by cellular protrusions of the blastoderm. The conspicuous large spines on the first blastodermic cuticle are formed by the evaginations of the blastoderm. Tendrils of the primary dorsal organ run between the first and second blastodermic cuticles.     

Embryogenesis of a springtail,Tomocerus ishibashii (Collembola,tomoceridae): External morphology

The external features of the developing embryos of the springtail, Tomocerus ishibashii, are described. The clypeolabral anlage arises as a single, unpaired swelling. The entognathy is completed by the ventral growth of the tergal anlagen of mandibular, maxillary, and labial segments. These anlagen also form the posterior part of the cranium. The palpi of maxilla and labium are homologous with the telopodites, and proximal parts of these head appendages are homologous with the coxopodites. The sternal element of the labial segment does not participate in the postmentum formation. The anlagen of abdominal appendages appear in the first to the fourth abdominal segments. The first, third, and fourth appendage anlagen form the ventral tube, tenaculum, and furcula, respectively. The fused proximal parts of the first, third, and fourth appendage anlagen are homologous with the coxopodites, and the distal parts which do not fuse are homologous with the telopodites. The anlagen of the second abdominal appendages become flattened and spread over the ventral side of this segment. The ventral structures of the first four abdominal segments are appendicular in origin.     

中国鳞虫兆属Tom ocerus四新种(弹尾目:鳞虫兆科)

本文记述了中国鳞虫兆属Ｔｏｍ ｏｃｅｒｕｓ４新种,即紫胸鳞虫兆Ｔｏｍ ｏｃｅｒｕｓ（Ｔｏｍ ｏｃｅｒｉｎａ） ｐｕｒ－ｐｕｒｉｔｈｏｒｕｓ, ｓｐ． ｎｏｖ．, （四川）; 白鳞虫兆Ｔｏｍｏｃｅｒｕｓ（Ｔｏｍ ｏｃｅｒｉｎａ） ｃａｌｃｅｕｓ, ｓｐ． ｎｏｖ．, （四川）;巨鳞虫兆Ｔｏｍｏｃｅｒｕｓ（ｓ．ｓｔｒ．）ｍ ａｘｉｍｕｓ,ｓｐ．ｎｏｖ．（四川）;小鳞虫兆Ｔｏｍ ｏｃｅｒｕｓ（ｓ．ｓｔｒ．）ｅｍ ｅｉｃｕｓ, ｓｐ． ｎｏｖ． （四川）。模式标本保存于绵阳经济技术高等专科学校农艺系昆虫标本室。１． 紫胸鳞虫兆Ｔｏｍ ｏｃｅｒｕｓ（Ｔｏｍｏｃｅｒｉｎａ） ｐｕｒｐｕｒｉｔｈｏｒｕｓ, 新种（图１）鉴别特征：本种与Ｔ．（Ｔｏｍ ｏｃｅｒｉｎａ） ｍ ｉｎｕｔｕｓＴｕｌｌｂｅｒｇ １８７６ 非常相似,但其下列特征可与后者相区别：弹器齿节刺的刺式为５／７,１,弹器端节间齿数为５～７,爪内齿数为１,１,１,握弹器体上刚毛数为５支。正模：♀,四川峨眉山,１９－ ＩＶ－ １９９５,刘永琴;副模：４♀♀２♂♂,同正模。２． 白鳞虫兆Ｔｏｍ ｏｃｅｒｕｓ （Ｔｏｍ ｏｃｅｒｉｎａ） ｃａｌｃｅｕｓ, 新种（图２）鉴别特征：本种与Ｔ．（Ｔｏｍｏｃｅｒｉ     

中国鳞属一新种(弹尾目:鳞科)

记述采自中国长白山的鳞虫兆属 Ｔｏｍ ｏｃｅｒｕｓ １新种：长白山鳞虫兆 Ｔ． ｃｈａｎｇｂａｉｓｈａｎｅｎｓｉｓ, ｓｐ． ｎｏｖ．。模式标本保存于北京自然博物馆。     

Cryptic diversity,diversification and vicariance in two species complexes of Tomocerus (Collembola,Tomoceridae) from China

Springtails (Collembola) are a group of arthropods that are found in terrestrial ecosystems throughout the world. Two species complexes, Tomocerus ocreatus and T. nigrus complexes, are widely distributed in the southern and northern parts of eastern China, respectively. There is a poor understanding of the species diversity within these complexes and of the factors affecting their diversification and dispersal. Species delimitation using a general mixed Yule coalescent model and a Bayesian multilocus approach recognized 22 DNA‐based species. This supports the presence of extensive cryptic diversity in species that are geographically widespread. In addition to genetic differences, we discovered corresponding morphological differences in jumping organs among the major clades. Analyses of divergence times and historical biogeographical processes revealed that ocreatus and nigrus complexes originated in southern and northern China, respectively. We estimated their divergence at 27.8–44.9 Mya during the Eocene–Oligocene, at the time when the transmeridional Qinling–Dabie Mountains uplifted and formed the north–south geographical boundary of eastern China. Diversification analyses suggest that the subsequent orogenesis of the Qinghai–Tibetan Plateau in western China had little impact on divergences within the two species complexes so that they maintained their geographical patterns from the Paleogene to the present day. Our findings also point to a potentially important influence of the Qinling–Dabie Mountains on patterns of animal speciation and distribution in China.     

Population structure, gene flow and evolutionary relationships in four species of the genera Tomocerus and Pogonognathellus (Collembola, Tomoceridae)

Genetic diversity and gene flow at 10 allozyme loci were investigated in 13 natural populations of four species of the collembolan genera Tomocerus and Pogonognathellus. Levels of observed heterozygosity were found to vary from 0.033 in P. longicornis to 0.120 in T. vulgaris. Average Nei's genetic distances (D) ranged from D = 0.222 between populations of P. ftavescens to D = 0.647 between populations of T. vulgaris. Genetic distances between species were always 1.000 and the highest value (D = 4.321) was between P. longicomis and one population of T. vulgaris. Values of F_sr were very high in all species (from 0.567 to 0.696) and levels of gene flow (Nm) derived from F_ST and the private allele method were low (Nm <1). Gene flow was significandy higher only in two subsets of populations of T. vulgaris. The Plio-Pleistocenic geological rearrangements and the effect of stochastic events, such as genetic drift, are invoked to explain the different levels of genetic divergence between and within species. Distance- and character-based approaches were used to reconstruct evolutionary relationships between and within species. While the monophyly of all species was confirmed, the results did not univocally support the monophyly of the two genera, leaving the question of their generic or subgeneric status unresolved.     

A peculiar cave species of Tomocerus (Collembola,Tomoceridae, Tomocerinae) from Vietnam,with a discussion of the postantennal organ and prelabral chaetae in Tomocerinae

The first Tomocerus species with a postantennal organ (PAO) in the adult stage is described from Vietnam. Tomocerus postantennalis sp. n. differs from the other PAO-possessing tomocerid, Tomolonus reductus Mills, 1948, mainly in the morphology of PAO, the number of ocelli, the number of chaetae in trochantero-femoral organ and several features of the furca. The new species is placed in Tomocerus because of the presence of a toothlet on the outer basal mucronal tooth and the absence of the diagnostic character states of Plutomurus Yosii, 1956 and Aphaenomurus Yosii, 1956. Besides the presence of PAO, the new species is peculiar in having six prelabral chaetae, instead of four as in other Tomocerus species. The new species is similar to Tomocerus folsomi Denis, 1929 and Tomocerus ocreatus Denis, 1948 in the type of dental spines but different from them in the body colour, the relative length of antennae to body, the number of unguis inner teeth and the number of mucronal intermediate teeth.     

The springtail (Hexapoda,Collembola) fauna of Wrangel Island

Analysis of the collembolan fauna of Wrangel Island based on published data and recent sampling was performed. The species richness of springtails on the island (89 species from 41 genera and 12 families) exceeds the levels characteristic of most other insular territories of the Arctic Ocean, except for Greenland and some small islands located within the southern tundra belt. The specificity of the species composition of the springtail fauna on the island depends on a high proportion of species with amphi-Pacific ranges, reflecting its formation within the Beringian Arctic sector. At the same time, the structure of the springtail fauna of the island is rather similar to those of the known tundra faunas of the Siberian sector, with no anomalies being detected.     

Resource allocation in Tomocerus flavescens (Insecta,Collembola): a study with C-14-labelled food

Summary Individuals of the collembolan species Tomocerus flavescens from a beech wood on limestone near Göttingen (West Germany) were fed with C-14-labelled algae in the laboratory. On an average, T. flavescens exchanged almost all of its endogenous carbon after 3.4 moults. In periods of starvation the interval between two ecdyses increased from 5.2 to 6.1 days (17°C). The assimilation efficiency was determined in three independent ways (carbon use of different algal components; A=C-FU; C=P+R+FU, A=P+R): it ranged from 0.34 to 0.40 (converted to energy values 0.30 to 0.45). It could be raised in phases of high mobility and in periods of food shortage by lowering the gut passage rate. The decrease in feeding activity (demonstrated by gut contents analyses) in case of a detoriation of the food quality and the food availability can be partly compensated in this way. For individuals which did not reproduce, the ratio of production to assimilation was 0.24 (converted to energy values 0.31). T. flavescens showed an increased body growth in connection with reaching sexual maturity. Specific demands of nutritive substances in periods of physiological changes could possibly be compensated by a more efficient resource use as well as by differences in resource allocation. The studied population of T. flavescens was compared both with that of a Danish beech forest and general data from the literature. The production of few eggs rich in energy, the high weight of newly hatched individuals, the high growth as well as the storage of carbon connected with egg production, the comparatively low respiration metabolism and the high amount of energy invested in search for food have to be regarded as an adaptation to the habitat. According to conservative estimates climbing individuals of this species consume 0.83 g of the algal dry mass growing on one beech tree during one summer. T. flavescens mainly uses the ethanol-soluble components of the algae.     

Redescription of Plutomurus kelasuricus Martynova, 1969 (Collembola: Tomoceridae) from Georgian caves

The collembolan species Plutomurus kelasuricus Martynova, 1969 Martynova, E. (1969): Springtails of the family Tomoceridae (Collembola) from the fauna of the USSR [in Russian]. Entomological Review, 43, 299–314. [Google Scholar] is redescribed and illustrated based on material sampled in Georgian caves. Differences from the morphologically similar P. abchasicus Martynova, 1969 Martynova, E. (1969): Springtails of the family Tomoceridae (Collembola) from the fauna of the USSR [in Russian]. Entomological Review, 43, 299–314. [Google Scholar] are discussed. A key to species of the genus Plutomurus found in the Caucasus is provided.     

Experimental studies of ice nucleation in an Antarctic springtail (Collembola, Isotomidae)

Ice nucleation was studied in field-fresh and acclimated (4 degrees C without food for 11-20 days) samples of the springtail Cryptopygus antarcticus Willem (Collembola, Isotomidae) at Rothera Research Station, Adelaide Island on the Antarctic Peninsula. Ice nucleator activity (INA) was measured by a freezing droplet technique in addition to supercooling point (SCP) profiles and polyol, sugar, and water contents. Field and acclimated samples showed bimodal SCP distributions with a distinct high group (HG; mean SCP -8 to -10 degrees C) and low group (LG: mean SCP -23 to -25 degrees C), which were significantly different. Acclimation at 4 degrees C increased the proportion of individuals in the LG relative to that in the HG without significant effects on the mean SCP of both groups. INA of the HG was significantly greater than that of the LG, and acclimation further reduced the INA of the LG. The number of active ice nucleator agents (INAs) calculated for the HG of field samples increased by 23-100 times over the temperature range -5 to -8 degrees C compared to only 7 times for the LG over the same range. These differences were accentuated in the acclimation experiments. Glucose and galactose were the main carbohydrates in both field and acclimated springtails, with the latter compound occurring in almost twice the concentration in the LG compared with that in the HG. Acclimation reduced the concentration of both compounds (glucose by 77% and galactose by 54%), whereas water content increased significantly. Digestion of food may have continued during acclimation at 4 degrees C, which could reduce the LG INA. Lowering of temperature over time is more likely to elicit a cold hardening response than constant temperature acclimation. INA numbers calculated at the nucleation temperatures for C. antarcticus samples were higher in the LG than in the HG. However, inactivation of INAs may be a key mechanism underlying cold hardening in this species, either by sequestration within the cellular matrix or by being only seasonally active.     

Formation of springtail (Collembola) communities during colonization of ant-hills

Formation of springtail assemblages in resettled nests of the ant Formica aquilonia was studied. The species composition and relative abundance of springtails was shown to depend on the age, hydrothermal regime, and condition of the ant nest. The abundance and species diversity of springtails increases during the initial period of the ant-hill growth. Old nests are characterized by the minimum species diversity, with one or two distinct dominants. High ecological plasticity, including resistance to high temperatures and low humidity, allows Lepidocyrtus violaceus to maintain its stable dominant position in the ant-hills. Species with relatively narrow hygro-and thermopreferenda quickly occupy temporal favorable niches, reach high abundance, and quickly disappear as the microclimatic conditions change.     

中国鳞属Tomocerus一新种(弹尾目:鳞科)

本文记述了中国鳞属1新种:王朗鳞Tomocerus(Tomocerina)wanglangensis,sp.nov.。模式标本保存于西南科技大学生命科学与工程学院昆虫标本室。王朗鳞虫兆Tomocerus(Tomocerina)wanglangensis,新种(图1~9)本新种和白鳞虫兆T.(Tomocerina)calceusLiuetal.1999非常相似,但新种爪内缘齿为1,1,1;弹器齿节刺简单、不具褶,刺序为2/1,Ⅰ,1,Ⅰ;弹器端节间齿数为2。正模♀,四川省平武县王朗,1800m,06-Ⅹ-2002,刘永琴;副模:3♀♀,5♂♂,同正模。     

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.     

The ultrastructure of the ejaculatory duct in the springtail Orchesella villosa (Geoffroy) (Hexapoda, Collembola) and the formation of the spermatophore

The initial part of the ejaculatory duct of Orchesella villosa contains a “valve” and a “sorter” avoiding respectively the reflow and allowing the separation of the secretion for the spermatophore stalk from the sperm fluid. For most of its length, the ejaculatory duct lumen is divided into two parts: in the dorsal part the sperm fluid flows while in the ventral district the secretion for the stalk occurs. Laterally, on both sides of the duct, longitudinal muscle fibers are present. The epithelium of the dorsal region consists of two types of long secretory cells; the most peculiar of them are those provided with extracellular cisterns flowing directly into the duct lumen as it occurs in 1st type of epidermal cells. These cells could be involved in the control of the viscosity of the sperm fluid. The second type of cells produce a secretion probably involved in the formation of the outer coat of the apical sperm droplet. The ventral epithelium consists of short cells contributing to the enrichment of the secretion for the spermatophore stalk and perhaps also to the viscosity of the secretion flowing in the lumen. In the distal part of the ejaculatory duct, the ventral district is provided with a thick layer of muscle fibers and with 3 + 3 cuticular laminae dividing the lumen into a series of slits through which the secretion of the stalk is squeezed out into filaments. This organization allows the twisting and hardening of these filaments. A drop of sperm fluid is laid on top of the long and rigid spermatophore stalk.     

The midgut of Tomocerus minor lubbock (Insecta,Collembola): Ultrastructure,cytochemistry, ageing and renewal during a moulting cycle

The midgut cells of Tomocerus minor (Insecta, Collembola) were examined with the electron microscope and cytochemically. The midgut epithelium consists of a series of cells characterised by numerous mineral concretions scattered throughout the cytoplasm. Mitochondria are abundant; microvilli are well developed at the apical surface of the cell. A zonula continua (continuous junction) characterises the apical contact region of these cells. Polysaccharides, glycoproteins and carbohydrate components have been demonstrated on the surface of microvilli. Peritrophic membranes surround the food bolus and preserve midgut cells from mechanical abrasion. Lysosomes are present during the alimentary period and show strong acid phosphatase activity. During an intermoulting cycle, two stages can be observed: (1) the postexuvial feeding period during which cytoplasmic extrusions appear at the apical part of the cell: lysosomes increase in number and autophagic vacuoles appear. (2) The preexuvial fasting period; a new epithelium grows beneath the old one and pushes it into the lumen. Degeneration processes can be observed in the old epithelium. This excretory reactivity of the midgut epithelium has been compared to the cycle of the cuticle.     

Mesoderm formation in the anuranXenopus laevis (Daudin)

Summary UsingXenopus blastulae of stage 9^–, recombinates were made of the animal, ectodermal cap (zones I.II) and the vegetative, endodermal yolk mass (zone IV) (see Fig. 1). For the experiments either the entire ectodermal cap (A.B), the single outer layer (A) or the stratified inner layer (B) were used.A comparison of the quantitative composition of the recombinates and the corresponding isolates—on the basis of absolute values expressed in units of section surface area—demonstrates unequivocally that the entire mesoderm originates from the ectodermal half of the anuran egg under an inductive influence emanating from the endodermal half. This holds for recombinates of the vegetative yolk mass with the entire ectodermal cap as well as with its outer or inner layer alone.A comparison of mesoderm formation in recombinates of the entire ectodermal cap or with its outer or inner layer with the vegetative yolk mass shows that in all cases mesoderm formation is proportional to the amount of ectoderm available. In addition, the outer layer of the ectoderm is partially endodermized which may be brought in relation with the fact that in normal development an endodermal lining extends upwards from the endodermal mass, which, among other things, covers the prechordal mesoderm on the outside.The outer layer of the ectoderm has markedly lower neural competence than the inner layer, from which in normal development the bulk of the neural material arises.     

A new relictual and highly troglomorphic species of Tomoceridae (Collembola) from a deep Croatian cave

Tritomurus velessp. n. (Tomoceridae) is described from a Croatian cave. It is characterized by troglomorphic features (absence of eyes, reduced pigmentation, slender claw, pointed tibiotarsal tenent hairs) that only compare, among Tomoceridae, to the microendemic species Tritomurus falcifer from the Pyrénées. Tritomurus veles also shares with Tritomurus falcifer the absence of macrochaetae on head, a presumably non-adaptive character that within Tomoceridae is unique to these two species. Both species have no known epigean relatives in their respective distribution areas and can be considered as relictual.     

New insight into the systematics of Tomoceridae (Hexapoda,Collembola) by integrating molecular and morphological evidence

Tomoceridae is common but among the most problematic groups of Collembola. Its position within Collembola and the relationships within the family remain obscure. This also extends to the generic division of the subfamily Tomocerinae that remains controversial. This study examines these issues by integrating both molecular and morphological evidence. Our molecular phylogeny based on rDNA sequences supports the monophyly of Tomoceridae and the sister relationship between Tomocerinae and Lepidophorellinae. Reconstructions and tree topology tests constraining monophyly did not resolve the relationships between Tomoceridae and other collembolan groups. We also examined the morphology of the first instar (primary) larvae, which has significant phylogenetic value among higher Collembola. Mapping primary chaetotaxy onto our molecular phylogeny provided further evidence for the unique position of Tomoceridae within Entomobryomorpha and Collembola. The monophyly and subfamilial classification within Tomoceridae were validated here, whereas its position among Collembola will need further studies in a broader consideration across the major collembolan orders. Within Tomocerinae, the monophyly of Pogonognathellus was demonstrated, but the status of Tomocerus and Tomocerina is still to be resolved.     

Ultrastructural and molecular identification of a new Rickettsia endosymbiont in the springtail Onychiurus sinensis (Hexapoda, Collembola)

In this paper we provide microscopic and molecular evidence for the presence of an endosymbiontic bacterium in male and female gonads of the soil arthropod Onychiurus sinensis. The sequence of the gene encoding for the 16S rRNA shows that the bacterium is a member of the genus Rickettsia, and some anomalies presumably associated with the presence of these microorganisms have been detected. Although the Rickettsia found in O. sinensis has the smallest genetic divergence with Rickettsia bellii, the phylogenetic analysis fails to find support for a sister-group relationship between these two species, rather suggesting that most Rickettsia species/strains isolated in various arthropods have rapidly evolved and diversified in what appears to be a sudden burst of evolution.