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.     

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。     

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.     

Mesoderm formation in a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The mesoderm formation of Tomocerus ishibashii (Collembola : Tomoceridae) is described. Mesodermal cells are formed after the beginning of the formation of the primary dorsal organ, and originate from the entire region of the embryonic area. After completion of the blastodermic cuticles, cells of mesoderm and ectoderm concentrate towards a ventral midline and form a well-defined 2-layered germ band. The manner of mesoderm formation in the Collembola is similar to that in Diplura and Myriapods, except for the Chilopoda; the mesoderm of the Thysanura s. lat. and Pterygota originates from a localized zone of the embryo. Within the Hexapoda, mesoderm formation is categorized into 2 types: Type 1—unlocalized origin, in the Collembola and Diplura, and Type 2—localized origin, in the Thysanura s. lat. and Pterygota. Types 1 and 2 are thought to be plesiomorphic and apomorphic, respectively.     

Endonura Cassagnau in Iran,with a key to species of the genus (Collembola,Neanuridae, Neanurinae)

Three new species of Endonura are described from Iran. Endonura dichaeta sp. n. can be recognized by an ogival labrum, head without chaetae O and E, chaeta D connected with tubercle Cl, tubercle Dl with five chaetae on head, absence of tubercles Di on thorax I and tubercle (Di+Di) of thorax V with 2+2 chaetae. Endonura ceratolabralis sp. n. is characterized by large body size, reduction of labral chaetotaxy, ogival labrum, head without chaeta O and fusion of tubercles Di and De on first thoracic segment. Endonura persica sp. n. is distinguished from its congeners by a nonogival labrum, absence of chaeta O, tubercles Dl and (L+So) with five and eight chaetae respectively and claw with inner tooth. The key to all species of the genus is given.     

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.     

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.     

Ritual behaviors associated with spermatophore transfer in Deuterosminthurus bicinctus (Collembola: Bourletiellidae)

The springtail Deuterosminthurus bicinctus, similarly to other members of Bourletiellidae, use their antennae, legs or heads to monopolize, stimulate, and direct female partners to spermatophores. The mating behavior of this species was examined by analysis of video recordings made on the leaves of its host plant. The characteristic stages of the behavioral sequence leading to sperm transfer were (1) preliminary courtship, highly variable in time and intensity—a male (rarely a female) endeavors to recruit a partner for further courtship, (2) push-and-retreat ritual—a rigid and rhythmical head-to-head dance of both partners, composed of some 180 phases and interspersed by several (3) free turns of a male and ended by (4) spermatophore deposition in front of a female, followed by stimulation of a female by a male to walk over the spermatophore, (5) spermatophore pick-up by a female with her gonopore, or alternately, (6) total spermatophore consumption by a female, as happened in 29% of the observed pairs. The mating ended always with (7) competition by both partners for the consumption of spermatophore residuals, usually won by the female. Comparison of mating elements and morphological features of partners engaged in mating sessions that ended either with spermatophore pick-up or consumption did not reveal any substantial differences. This suggests that changes in female motivation (hunger, state of impregnation) may be crucial for the male success in sperm transfer to a female. A digital video image is available at An erratum to this article can be found at     

New species of Phallodrilus (Oligochaeta,Tubificidae) from caves of northern Spain and southwestern France

A taxonomic account of a collection of Phallodrilus species inhabiting caves is given. The following three new species are described: P. subterraneus, P. crypticus and P. labouichensis. New material of P. aquaedulcis Hrabe, 1960 from northern Spain and southwestern France is reported. This species was previously known from West Germany. The relationship between Phallodrilus cave species and littoral and deep-sea species is discussed.     

A new species of Erebonectes (Copepoda,Calanoida) from marine caves on Caicos Islands,West Indies

The insects of a small and cold aestival stream in northern Sweden were studied from semiquantitative kick-samples taken every ten days throughout the ice-free season in 1987. The stream froze solid at least in some sections in some winters. In addition to dipterans that were only identified to the family level, five ephemeropteran, four plecopteran, 11 coleopteran (Dytiscidae), and four trichopteran (Limnephilidae) species were found. Baetis bundyae Lemkuhl, Heptagenia orbiticola Kluge (Ephemeroptera), Nemoura cinerea (Retzius) (Plecoptera), Hydroporus palustris (L), Agabus setulosus (J. Sahlberg) (Coleoptera), Anabolia concentrica (Zetterstedt), Limnephilus coenosus Curtis, and Micopterna sequax McLachlan (Trichoptera) were all common. Malaise trap data are given for Plecoptera. The two ephemeropteran species Heptagenia orbiticola and Metretopus alter Bengtsson, and the plecopteran species Nemoura viki Lillehammer were recorded for the first time in Sweden. The remarkable species composition was attributed to the harsh abiotic and poorly studied environment. We predicted that the larval composition of this stream was representative for aestival streams in northernmost Fennoscandia.     

Growth and reproduction of fungal feeding Collembola as affected by fungal species,melanin and mixed diets

Fungal feeding soil invertebrates feed on a wide spectrum of fungal species suggesting that mixed diets increase fitness. We investigated relationships between food preferences for seven saprophytic fungal species/forms and fitness parameters (mortality, growth, time to reproduction, reproduction, egg size) in two Collembola species, Folsomia candida and Protaphorura armata. The fungal species/forms studied included the wild type and a melanin-deficient form of Aspergillus fumigatus to investigate the role of melanin in collembolan nutrition. Also, three mixed diets consisting of a preferred fungal species (Cladosporium cladosporioides) and species of intermediate or low food quality were investigated. Both Collembola species preferred similar fungal species/forms as food. Food preference generally matched fitness parameters, i.e. growth and reproduction of Collembola was at a maximum when feeding on preferred fungi. This was not the case for A. fumigatus. The wild type and the melanin-deficient form ranked among the least preferred fungi. Growth and reproduction of Collembola were low when feeding on the wild type but high when feeding on the melanin-deficient form indicating that the Collembola misjudged the food quality of the latter in the preference tests. The results show for the first time that genes driving melanin syntheses (pksP) strongly affect the food quality of fungi for fungal feeding invertebrates. Feeding on mixed diets generally increased growth and reproduction of Collembola except when the diets included toxic species (Penicillium sp.). The results support the nutrient balance hypothesis and also show that the detection of toxic species in the diet is important. They indicate that the widespread generalist feeding mode of Collembola maximizes fitness if toxic fungal species are avoided. The fitness parameters growth, reproduction and time until onset of reproduction were correlated closely but egg volume, which also varied with fungal diet, correlated poorly with the other fitness parameters. Variation in egg size with fungal diet shows that the diet of Collembola may have transgenerational effects.     

Revisiting Lepidonella Yosii (Collembola: Paronellidae): character overview,checklist of world species and reassessment of Pseudoparonella doveri Carpenter

Characters used in the taxonomy of the genus Lepidonella Yosii, 1960 (Collembola: Paronellidae) are listed and discussed. Several new ones are introduced. An overview of pseudopore patterns across Collembola is presented, with several new locations of these structures across Entomobryomorpha. Their interest at different taxonomic level is underlined. The genus Lepidonella is redefined. The American species L. marimuti Soto Adames &; Bellini, 2015 Soto-Adames FN, Bellini BC. 2015. Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea). Florida Entomologist. 98(1):330–341.[Crossref], [Web of Science ®] , [Google Scholar] is placed in incertae sedis among Lepidonella. Lepidonella species of the world are listed with synonymies and combinations. The Malaysian troglobitic species Pseudoparonella doveri Carpenter, 1933 Carpenter GH. 1933. XIX. Fauna of the Batu caves, Selangor. Journal of the Federated Malay States Museum. 17:217–221. [Google Scholar] is redescribed in detail, with emphasis on its pattern of antennal chaetae, and transferred to the genus Lepidonella. Its close similarity with L. lecongkieti Deharveng &; Bedos, 1995 Deharveng L, Bedos A. 1995. Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae). Bulletin de la Société entomologique de France. 100(1):21–24. [Google Scholar] from southwestern Vietnam caves is underlined. This disjunct distribution is briefly discussed.     

贵州部分洞穴裸灶螽的鉴定及其遗传多样性分析

贵州是中国具有丰富喀斯特洞穴生态系统的省份之一,裸灶螽作为洞穴中的优势物种又为生态环境指示种,其遗传多样性能够反应洞穴生物多样性.为评估洞穴开发对洞穴生物多样性的影响,本研究通过形态鉴定和分子辅助鉴定的方式对来自贵州省毕节、黔西南、铜仁和黔南4个地区8个洞穴97号裸灶螽标本进行鉴定,并对其遗传多样性进行分析.结果共鉴定...     

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.     

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.