Research Article |
Corresponding author: Spyros Sfendourakis ( sfendourakis.spyros@ucy.ac.cy ) Academic editor: Elisabeth Hornung
© 2015 Spyros Sfendourakis, Stefano Taiti.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sfenthourakis S, Taiti S (2015) Patterns of taxonomic diversity among terrestrial isopods. In: Taiti S, Hornung E, Štrus J, Bouchon D (Eds) Trends in Terrestrial Isopod Biology. ZooKeys 515: 13–25. https://doi.org/10.3897/zookeys.515.9332
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The publication of the world catalog of terrestrial isopods some ten years ago by Schmalfuss has facilitated research on isopod diversity patterns at a global scale. Furthermore, even though we still lack a comprehensive and robust phylogeny of Oniscidea, we do have some useful approaches to phylogenetic relationships among major clades which can offer additional insights into isopod evolutionary dynamics. Taxonomic diversity is one of many approaches to biodiversity and, despite its sensitiveness to biases in taxonomic practice, has proved useful in exploring diversification dynamics of various taxa. In the present work, we attempt an analysis of taxonomic diversity patterns among Oniscidea based on an updated world list of species containing 3,710 species belonging to 527 genera and 37 families (data till April 2014). The analysis explores species diversity at the genus and family level, as well as the relationships between species per genera, species per families, and genera per families. In addition, we consider the structure of isopod taxonomic system under the fractal perspective that has been proposed as a measure of a taxon’s diversification. Finally, we check whether there is any phylogenetic signal behind taxonomic diversity patterns. The results can be useful in a more detailed elaboration of Oniscidea systematics.
Bodiversity, diversification, systematics, fractals, phylogeny, species richness, taxonomic asymmetry
Terrestrial isopods constitute one of the most remarkable lineages of invertebrates that managed to conquer land. Modern species represent almost all evolutionary steps that enabled them to leave the marine environment and occupy almost the whole range of terrestrial habitat types. This makes them a unique case within global biodiversity and offers lots of opportunities to biological research, especially in fields like evolution, ecology and ecophysiology (
Terrestrial isopods, the suborder Oniscidea within the order Isopoda, are currently considered to be a monophyletic taxon even though their monophyletic origin has been questioned in the past (see
From an ecological point of view, Oniscidea live in almost all biomes, having successfully invaded most areas of the world, with the exception of the poles and very high elevations (>4,800 m,
Oniscidea probably originated in the Carboniferous (
The study of taxonomic diversity aims to explore such patterns across different taxonomic levels. Despite its sensitiveness to biases in taxonomic practice it has proved useful in exploring diversification dynamics in characteristic biota (e.g.,
In the present work we attempt an analysis of taxonomic diversity patterns among Oniscidea based on an updated world list of species, exploring species diversity at the genus and family level, as well as the relationships between species per genera, species per families, and genera per families. Even though the assignment of genus and family status for a group of species or clades is arbitrary, experts in each higher taxon usually follow a similar approach, so that the study of such patterns is still meaningful to a considerable extent. Also, we do know that several families (or even genera) of Oniscidea might not be monophyletic, and this could lead to uncertainties in results from such a taxonomic diversity analysis. Nevertheless, this kind of analysis actually helps towards identifying such problems. For example, the exploration of a possible fractal structure in the isopod taxonomic system, which has been proposed as a measure of diversification, is a useful tool for this, and we do address the issue herein. Finally, we check whether there is any phylogenetic signal behind taxonomic diversity patterns.
In order to compile a complete species list of valid isopod species, we used as a basis the world catalog published by
Species description rates were calculated from dates appearing in current nomenclature.
For phylogenetic information, we used the morphological analysis by
In order to test for skewness in the frequency distribution of species richness at different taxonomic levels we used the standardized skewness metric and the Shapiro Wilk test for deviation from normality. In all other analyses we applied standard linear regressions and the Pearson product moment, or the Spearman rank correlation coefficient.
In the ten years after the publication of the electronic version of the world catalog of terrestrial isopod species (
The list of families with the respective numbers of genera and species is given in Table
List of families with their respective numbers of genera and species, the latter separately for those in known genera and those of uncertain generic assignment.
Family | Number of genera | Species in known genera | Species of uncertain generic assignment | Total |
---|---|---|---|---|
Armadillidae | 80 | 579 | 118 | 697 |
Philosciidae | 107 | 501 | 36 | 537 |
Trichoniscidae | 87 | 492 | 2 | 494 |
Porcellionidae | 19 | 326 | 7 | 333 |
Armadillidiidae | 14 | 256 | 0 | 256 |
Eubelidae | 50 | 253 | 2 | 255 |
Agnaridae | 14 | 157 | 10 | 167 |
Platyarthridae | 7 | 122 | 0 | 122 |
Trachelipodidae | 6 | 109 | 4 | 113 |
Scleropactidae | 26 | 107 | 0 | 107 |
Ligiidae | 6 | 95 | 0 | 95 |
Styloniscidae | 10 | 81 | 1 | 82 |
Cylisticidae | 5 | 66 | 0 | 66 |
Detonidae | 4 | 39 | 0 | 39 |
Halophilosciidae | 3 | 35 | 0 | 35 |
Oniscidae | 5 | 31 | 10 | 41 |
Alloniscidae | 2 | 23 | 2 | 25 |
Tylidae | 2 | 22 | 0 | 22 |
Spelaeoniscidae | 7 | 20 | 0 | 20 |
Delatorreidae | 3 | 18 | 0 | 18 |
Dubioniscidae | 3 | 15 | 0 | 15 |
Rhyscotidae | 2 | 13 | 0 | 13 |
Olibrinidae | 4 | 11 | 0 | 11 |
Scyphacidae | 2 | 11 | 0 | 11 |
Bathytropidae | 1 | 10 | 0 | 10 |
Balloniscidae | 2 | 8 | 0 | 8 |
Titaniidae | 5 | 6 | 0 | 6 |
Tendosphaeridae | 3 | 4 | 0 | 4 |
Stenoniscidae | 2 | 4 | 0 | 4 |
Pudeoniscidae | 2 | 4 | 0 | 4 |
Irmaosidae | 1 | 2 | 0 | 2 |
Mesoniscidae | 1 | 2 | 0 | 2 |
Schoebliidae | 1 | 2 | 0 | 2 |
Berytoniscidae | 1 | 1 | 0 | 1 |
Bisilvestriidae | 1 | 1 | 0 | 1 |
Hekelidae | 1 | 1 | 0 | 1 |
Turanoniscidae | 1 | 1 | 0 | 1 |
Unknown | 37 | 90 | 0 | 90 |
Total | 527 | 3,518 | 192 | 3,710 |
The family with the highest species richness is Armadillidae, followed by Philosciidae and Trichoniscidae. The same three families are also the richest in genera, albeit in a different order, with Philosciidae first, followed by Trichoniscidae and Armadillidae. There are seven monogeneric families, four of them with monotypic genera. Species richness is significantly correlated with genera richness (Spearman rank correlation coefficient: rs = 0.91, p < 0.001).
Species descriptions per decade showed a bimodal distribution in the last century with most of the currently valid species being described either in the first half of the 20th century, especially in the ‘20s, or from 1960 to 1990 (Fig.
Frequency distributions of isopod richness are significantly right-skewed (for genera within families: skewness = 6.82, Shapiro-Wilk test p < 0.001; for species within genera: skewness = 62.3, Shapiro-Wilk test p < 0.001; for species within families: skewness = 5.7, Shapiro-Wilk test p < 0.001). This means that most families and genera consist of few genera and species, respectively, while very rich lineages are rare (Fig.
The number of species per genus in a family is not predicted by the number of genera per family (Fig.
The frequency of genera is negatively correlated with the number of species per genus (in logarithmic space: r = -0.88, p < 0.001; Fig.
Linear regression of the frequency of genera (fGenera) against their respective species richness, revealing the fractal nature of terrestrial isopod taxonomy. The slope of the regression (1.14) gives the fractal dimension. Unit frequency values – genera with unique values of species richness – have been excluded in order to avoid a long queue of zeros that smoothens the slope only due to the fact that the size of large genera is more probable to be unique.
Species richness values per family were mapped on the available phylogenetic trees for Oniscidea (Fig.
Terrestrial isopods are the largest suborder of Isopoda and actually the only group of Crustacea that has managed to exploit almost the whole range of terrestrial ecosystems. The ca. 3,700 species known so far include clades that have evolved a variety of morphological, physiological and behavioral characters offering unique solutions to key problems pertaining to the adaptation to the life on land, so that today they represent almost all transitional stages from marine to extremely arid environments.
According to the rate of species descriptions presented herein one might assume that the vast bulk of the global oniscid diversity has been known, and the total richness will not change to a significant degree in the near future. Nevertheless, we should note that the ‘plateau’ in the accumulated species richness observed in the last two decades might be better attributed to the decline in taxonomic expertise on the group. Indeed, there are very few active taxonomists of Oniscidea today. A large part of the world remains unexplored, especially the tropics, and the current trends in funding and ‘academic prestige’ do not leave much space for optimism that they will be explored soon. It is equally important to note that many thousands of caves around the world are expected to host hundreds, if not thousands, of isopod species, taking into account that Oniscidea are amongst the richest animal taxa in troglobitic species, most of which occur in one or a few local caves and/or other subterranean habitats. Furthermore, several analyses based on molecular markers reveal an even higher diversity among isopod taxa (e.g.,
The analysis of global diversity conducted herein reveals a strong right-skewed frequency distribution, so that Oniscidea mostly contain genera with few species and families with few genera. This is a pattern observed also in other animal taxa (e.g., hexapoda:
The correlation between number of species and number of genera in a family, in combination with the fact that numbers of species per genus cannot predict generic richness in a family, underlines the wide variation inside Oniscidea. This is because in addition to the somewhat trivial fact that many small genera – even monotypic – may be found in large families, the pattern is also based on the occurrence of very diverse genera in small families.
If fractal geometry of taxonomic systems indeed reflects real patterns of evolutionary diversification, then isopod diversity appears underestimated. The ‘fractal dimension’ of most arthropods and North American isopods (including freshwater and marine) is 1.50 according to
An intriguing question refers to the role of ‘key innovations’ in adaptive radiations, which would lead to prominent radiations in clades that have acquired some new feature offering significant selective advantages. Such a pattern would lead to very asymmetric phylogenies and key innovations could be mapped as defining synapomorphies of prolific clades. If the phylogeny of
It is absolutely necessary to have a robust phylogeny of Oniscidea families (and genera) in order to gain crucial insights into the evolution of this fascinating taxon. New techniques using Next Generation Sequencing can facilitate this task and provide very useful information.
We would like to thank the two anonymous reviewers for their thoughtful comments on the manuscript that helped us to improve the final text.