Abstract

BackgroundTowards realistic estimations of the diversity of marine animals, tiny meiofaunal species usually are underrepresented. Since the biological species concept is hardly applicable on exotic and elusive animals, it is even more important to apply a morphospecies concept on the best level of information possible, using accurate and efficient methodology such as 3D modelling from histological sections. Molecular approaches such as sequence analyses may reveal further, cryptic species. This is the first case study on meiofaunal gastropods to test diversity estimations from traditional taxonomy against results from modern microanatomical methodology and molecular systematics.ResultsThe examined meiofaunal Pseudunela specimens from several Indo-Pacific islands cannot be distinguished by external features. Their 3D microanatomy shows differences in the organ systems and allows for taxonomic separation in some cases. Additional molecular analyses based on partial mitochondrial cytochrome c oxidase subunit I (COI) and 16S rRNA markers revealed considerable genetic structure that is largely congruent with anatomical or geographical patterns. Two new species (Pseudunela viatoris and P. marteli spp. nov.) are formally described integrating morphological and genetic analyses. Phylogenetic analysis using partial 16S rRNA, COI and the nuclear 18S rRNA markers shows a clade of Pseudunelidae species as the sister group to limnic Acochlidiidae. Within Pseudunela, two subtypes of complex excretory systems occur. A complex kidney already evolved in the ancestor of Hedylopsacea. Several habitat shifts occurred during hedylopsacean evolution.ConclusionsCryptic species occur in tropical meiofaunal Pseudunela gastropods, and likely in other meiofaunal groups with poor dispersal abilities, boosting current diversity estimations. Only a combined 3D microanatomical and molecular approach revealed actual species diversity within Pseudunela reliably. Such integrative methods are recommended for all taxonomic approaches and biodiversity surveys on soft-bodied and small-sized invertebrates. With increasing taxon sampling and details studied, the evolution of acochlidian panpulmonates is even more complex than expected.

Highlights

  • The study of cryptic species, i.e. two or more distinct species classified as a single species due to the lack of morphological differences, augmented during the last 20 years [1]

  • Exceptions are members of taxa living in the mesopsammon which generally are assumed to be direct developers [9] or, as in case of acochlidian panpulmonate gastropods, may have planktonic larvae which remain in the interstitial spaces [10]

  • Until now, applying morphospecies concepts on tiny meiofaunal gastropods has never been tested by molecular analyses

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Summary

Introduction

The study of cryptic species, i.e. two or more distinct species classified as a single species due to the lack of morphological differences, augmented during the last 20 years [1]. Based on morphology, only 28 valid species, 20 of them mesopsammic, were described globally Several of these species such as Microhedyle remanei (Marcus, 1953) were considered to be widespread throughout Western Atlantic warm water sands, i.e. in Brazil, Colombia and Bermuda [11,12,13,14], and Pseudunela cornuta (Challis, 1970) was recorded to occur on the Solomon Islands (Melanesia) and near Hong Kong (South China Sea) [15,16]. Since the biological species concept is hardly applicable on exotic and elusive animals, it is even more important to apply a morphospecies concept on the best level of information possible, using accurate and efficient methodology such as 3D modelling from histological sections Molecular approaches such as sequence analyses may reveal further, cryptic species. This is the first case study on meiofaunal gastropods to test diversity estimations from traditional taxonomy against results from modern microanatomical methodology and molecular systematics

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