Collembola, the biological species concept and the underestimation of global species richness

Despite its ancient origin, global distribution and abundance in nearly all habitats, the class Collembola is comprised of only 8000 described species and is estimated to number no more than 50 000. Many morphologically defined species have broad geographical ranges that span continents, and recent molecular work has revealed high genetic diversity within species. However, the evolutionary significance of this genetic diversity is unknown. In this study, we sample five morphological species of the globally distributed genus Lepidocyrtus from 14 Panamanian sampling sites to characterize genetic diversity and test morphospecies against the biological species concept. Mitochondrial and nuclear DNA sequence data were analysed and a total of 58 molecular lineages revealed. Deep lineage diversification was recovered, with 30 molecular lineages estimated to have established more than 10 million years ago, and the origin almost all contemporary lineages preceding the onset of the Pleistocene (~2 Mya). Thirty-four lineages were sampled in sympatry revealing unambiguous cosegregation of mitochondrial and nuclear DNA sequence variation, consistent with biological species. Species richness within the class Collembola and the geographical structure of this diversity are substantially misrepresented components of terrestrial animal biodiversity. We speculate that global species richness of Collembola could be at least an order of magnitude greater than a previous estimate of 50 000 species

Further acquisitions on systematics relationships within the genus Orchesella (Collembola, Entomobryidae) using allozymes.

Inter- and intraspecific genetic variability has been studied in 17 populations of six different species of the genus Orchesella from central Italy by means of allozyme electrophoresis. The electrophoretic results are consistent with the determination of the species based on their patterns of pigmentation, revealing a very high level of genetic divergence between them as a result of the age of the genus and its long-time evolution from the mainland with populations from the Tuscan archipelago. Two different approaches for evaluating levels of interpopulational gene flow have revealed the presence of a remarkable genetic isolation between them. The reason for this differentiation has to be found in the paleogeographical rearrangements that occurred in the Mediterranean and the Apennine areas during the Tertiary and Quaternary

The ultrastructure of the spermathecae in the Collembola Symphypleona (Hexapoda)

The fine structure of the paired spermathecae of two Symphypleona Collembola, Sminthurus viridis and Allacma fusca, was studied. The spermathecal epithelium is rich in pigment granules, while secretory organelles are scarce, consisting of a few mitochondria and microtubules, scattered cisterns of endoplasmic reticulum and Golgi systems. Small isolated secretory granules are visible in the apical cell region, where irregular microvilli are present. Beneath the epithelium, large cells rich in endoplasmic reticulum and Golgi systems are present; they are separated from the epithelial cells by a thin basal lamina. The epithelium is lined by a thin cuticle, which forms thin extensions at the beginning of the spermathecal duct. At this level, muscle fibers are visible. The spermathecal lumen is filled with many spermatozoa. At molting, the spermathecal epithelial cells detach from the cuticle and a large electrontransparent space is evident between the cytoplasm and the cuticle. The cytoplasm has few pigment granules and the spermathecal lumen contains degenerating spermatozoa. After 3 days, the spermathecal epithelium regains its usual appearance and the spermathecal lumen is empty. The two species of Collembola Symphypleona do not have accessory glands. Compared with the Arthropleona Orchesella villosa, which has a single spermatheca and two accessory glands (Dallai et al. [2008] J Morphol 269:464–478), a great difference has occurred in the organization of the female genital system of these species. The results are discussed in light of the implications for reproductive behavior and egg protection in Symphypleona

Different sperm number in the spermatophores of Orchesella villosa (Geoffroy) (Entomobryidae) and Allacma fusca (L.) (Sminthuridae)

Members of the basal hexapod group Collembola perform indirect sperm transfer; males deposit spermatophores on the soil and these are then picked up by females for insemination. Orchesella villosa exhibits a dissociated mating behavior, while Allacma fusca has either associated mating behavior or, more commonly, produces spermatophores without pairing. Sperm ultrastructure in the two species has been studied by TEM and SEM observations to estimate the volume of these cells. The sperm number into each spermatophore has been determined by counting the .fluorescent nuclei after Hoechst staining of sperm droplets squash preparations. Sperm droplets at the apex of spermatophores contain numerous spermatozoa of typical appearance, wound up and with a long anterior appendage (peduncle). Sperm droplets of Orchesella villosa are smaller (about 10 times) than in A. fusca, however they contain substantially more sperm cells (about three times). These differences could be explained as follows: (a) sperm of O. villosa are shorter than those of A. fusca (58 and 107 mm, respectively) and thus the volume of a single sperm of O. villosa is about 10 times smaller than in A. fusca; (b) considering the volume of sperm droplets and of sperm cells, the estimated number of sperm cells would be higher in A. fusca than in O. villosa; and (c) the conspicuous reduction of the sperm cells observed in A. fusca is likely linked to the aberrant meiosis present in this species, as well as in all Symphypleona studied so far, that causes a loss of 50% of their sperm

The spermatogenesis and oogenesis of the springtail Podura aquatica Linné, 1758 (Hexapoda: Collembola)

Podura aquatica is a springtail of uncertain systematic position. Our study dealing with the ultrastructure of the spermatogenesis and oogenesis of this species is a contribution to a better knowledge of both the reproduction and the systematics of the taxon. In the male, the spermatogenesis proceeds in a similar way to that of other Collembola. Primary spermatocytes do not show synaptonemal complexes which, instead, are found in primary oocytes. Thus a genomic recombination seems to be present only in females, as it occurs in other springtails. Degeneration of secondary spermatocytes, as reported in some families of the Symphypleona, was not observed in P. aquatica. At the end of spermiogenesis, a rolled up sperm cell provided with an anterior long appendage adhering to the acrosome is produced. In the female, the oogenesis also proceeds in a conventional way with the production of eggs rich in yolk. A branched spermatheca is present at the end of the common oviduct, close to the genital opening. It contains many sperm in its lumen. Contrary to the globular appearance of sperm cells in the male genital ducts, in the spermatheca they are straight, elongated, and lack the long anterior appendage. P. aquatica shows a spermatogenesis, sperm structure, and oogenesis similar to those of other Collembola. In agreement with the results of recent phylogenetic studies, we confirm that P. aquatica is a member of Poduridae, and it does not belong to a group close to the Symphypleona

Comparative analysis of two edaphic zoocenoses (Acari Oribatida; Hexapoda Collembola) in the area of Orio al Serio Airport (Bergamo, northern Italy)

Species richness, abundance and diversity of oribatid mites and springtails in soil litter were investigated at four sites in the area of Orio al Serio Airport (Bergamo, Italy). A total of 60 oribatid mite and 35 springtail species were collected from the sites. The general low diversity and eveness along with high dominance may be related to the sensitivity of these edaphic zoocenoses to soil impoverishment which has been occurring in this part of the italian pre-Alps for many years.Quantitative and qualitative differences were found among the investigated areas. Results show that less frequently disturbed soils are capable of restoring oribatid and springtail population, leading to greater equilibrium. Results show that less frequently disturbed soils allow the restoration of more equilibrated oribatid and springtail populations. Species were divided into 8 autoecological and 11 chorological categories, Correspondance Analysis arranged these categories among the sites, confirming the above-mentioned trend

High levels of genetic variability and population differentiation in Gressittacantha terranova (Collembola, Hexapoda) from Victoria Land, Antarctica

Allozyme electrophoresis was used to asses genetic variability and differentiation in 22 populations of Gressittacantha terranova Wise (Hexapoda, Collembola) from a coastal area of Victoria Land between the Mariner Glacier and the Nansen Ice Sheet. Allelic frequencies were determined at five enzyme loci: Phi, Pgm, Hk, Mpi and Mdh. Levels of variability, estimated as rates of heterozygosity, were higher than those calculated for the same loci in taxonomically related and non-related species of non-Antarctic Collembola. Thus, in spite of the ecological simplicity of Antarctic terrestrial ecosystems, G. terranova is characterized by high levels of genetic variability, and the 22 populations could be divided into three geographic groups, separated by the Aviator and Campbell glaciers. Genetic differentiation reflects the geographic arrangment of the populations, suggesting that the glaciers are effective barriers to gene flow, and that the patchy distribution of collembolan species in Antarctica has the potential to induce, in the long term, microspeciation processes. Interestingly, detectable genetic differentiation was observed between six populations collected at Edmonson Point, even though these are very close to each other, indicating the impact of geographic isolation even within short distances. The only exception to the congruence between genetic and geographic structuring was provided by the population of Apostrophe Island, for which a recent introduction with individuals coming from southern populations is suggested