Aim
The genetic diversity and distribution of earthworm species in the Mediterranean Basin has been influenced by their low vagility, as well as by the basin's complex geological and climatic history. Within this context, our objective was to evaluate the evolutionary history of hormogastrid earthworms by exploring their phylogeny, reconstructing ancestral areas and identifying potential vicariance and dispersal events.
Location
Western Mediterranean region, encompassing the geographical range of the family Hormogastridae (four genera, 30 species and subspecies).
Methods
We sampled the easternmost area of the hormogastrid range and integrated 606 new sequences with all the molecular data available from previous studies on the westernmost area. Mitochondrial and nuclear markers were amplified and sequenced, and the hormogastrid phylogeny (using Bayesian and likelihood methods) and networks were investigated. Ancestral‐area reconstructions were implemented in rasp.
Results
Ancestral‐area reconstructions provided similar results for alternative phylogenetic hypotheses, placing the origin of Hormogastridae between southern France and the north‐eastern Iberian Peninsula, and highlighting north–central Sardinia as a key ancestral area for diversification of eastern Hormogastridae (Hormogaster redii and H. samnitica). Multiple vicariance and dispersal events were detected, indicating a complex evolutionary history. Sardinian populations of H. samnitica and H. redii were not monophyletic but were related to populations from other areas, suggesting that Sardinia is a more complex biogeographical region than anticipated. Our results identified a phylogenetic relationship between the southern Sardinian fauna and that of Sicily, as well as relationships between the fauna of north‐eastern Sardinia and those of the Italian Peninsula and the Tuscan Archipelago.
Main conclusions
Our results suggest a complex evolutionary history for hormogastrid earthworms, showing the possibility of many dispersal and vicariance events and multiple faunal interchanges between land‐masses, which may reflect the biogeographical complexity of the Mediterranean Basin.
The problem of reconciling earthworm taxonomy and phylogeny has shown advances with the application of molecular techniques, yet they have proven insufficient. Integrative systematics could solve this by combining multiple sources of evolutionary information. Relatively low diversity, restricted range and low nomenclatural conflict make Hormogastridae Michaelsen, 1900 a desirable target for an integrative systematics approach. The main systematic conflicts within this family are the polyphyly of the species Hormogaster pretiosa Michaelsen, 1899, the widespread presence of cryptic lineages, the lack of resolution of supraspecific relationships and the paraphyly of the genus Hormogaster Rosa, 1877 (found to be composed of four well-supported genus-level clades by molecular phylogenetic inference). This work integrates all the existing information by performing phylogenetic inference based on morphological, molecular and total evidence datasets, comparing their performance with the topology obtained by phylogenomic analyses. It also includes a comparative study of representatives of the main clades based on microcomputed tomography (µCT) reconstructions. The addition of morphological characters improved the resolution of the Hormogastridae tree; ancestral state reconstruction displayed the evolution of character states and provided morphological diagnoses for the genera within a new system, which incorporates information about ecological niches and biogeography.
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