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New material from the Miocene of Italy allows revision of the spatangoid genus Mariania Airaghi, 1901 and proposal of an emended diagnosis. Particularly characteristic, previously overlooked features of the genus include the presence of well-developed phyllodes made up from short, almost equidimensional plates in oral ambulacra II, III and IV. Unlike in other Spatangoidea, where the adoral plates rapidly become elongated towards the margin, they stay short in Mariania and are not constricted halfway between the peristome and the margin. In addition, most species of Mariania possess a characteristic domal profile with steep sides and lack a raised keel in aboral interambulacrum 5. Their petals are wide, open distally and extend almost to the margin. The plastron is not indented behind the episternal plates and the labral plate extends to the second ambulacral plates. Fascioles are missing in all specimens examined. The combination of these morphological features enable the separation of Mariania from the genera Macropneustes, Hypsopatagus and Spatangus, to which members of the genus have been assigned by previous authors. Cladistic analysis carried out to unravel the uncertain systematic position of Mariania failed to find well-supported relationships, but firmly places Mariania within the Brissidina. Most previous family attributions could be, however, ruled out. Based on the available data a placement within Spatangoidea seems most likely, where it takes up an intermediate position between maretiids, loveniids and spatangids. Three different species are identified within the studied sample: Mariania marmorae, the type species of the genus; M. stefaninii sp. nov. from the late Burdigalian-early Langhian of northern Italy; M. comaschicariae sp. nov. from the Burdigalian of Sardinia. These new species are distinguished from M. marmorae by their lower tests, shorter labral plates and shorter petals. Mariania comaschicariae sp. nov. differs from M. stefaninii sp. nov. by its lower test, more anterior apical disc and less numerous plates in the oral anterior paired ambulacra. Test morphology and parent rock sedimentology suggest that Mariania was an epifaunal echinoid, which lived in inner shelf environments, characterized by soft bottoms and a tropical climate. •

Spatangoids are probably the least resolved group within echinoids, with known topological incongruencies between phylogenies derived from molecular (very scarce) and morphological data. The present work, based on the analysis of 270 specimens of Spatangidae (Echinoidea, Spatangoida) trawled in the Sardinian seas (Western Mediterranean), allowed us to verify the constancy of some characters that we consider to be diagnostic at the genus level —such as the path of the subanal fasciole and the relationship between labrum and adjacent ambulacral plates —and to distinguish two distinct forms within the studied material. Based on morphological characters, morphometrics, and molecular analyses (sequencing of two mitochondrial markers: cytochrome c oxidase subunit1 (COI) and 16S), most of the individuals were classified as morphotype A and attributed to the species Spatangus purpureus, the most common spatangoid in the Mediterranean Sea, while a few corresponded to a different morphotype (B), genetically close to the species Spatangus raschi. Preliminary morphological analyses seemed to indicate that morphotype B specimens from Sardinia are slightly different from S. raschi and from Spatangus subinermis individuals, the second species of the family known to occur in the Mediterranean Sea. On the basis of morpho-structural observations and molecular analyses, comparing Mediterranean living forms with species from other areas (Central Eastern Atlantic, North Sea and neighboring basins, South African Sea, Philippines and Indonesian Archipelago, New Zealand, and Hawaiian Islands), the clear distinction of S. purpureus from several other species classified as Spatangus was confirmed. Based on the morphological and genetic differences, we propose to maintain the genus Spatangus including in it only the type species S. purpureus among the living species and to establish the new genus Propespatagus nov. gen. to include several other species previously classified as Spatangus. The clear distinction among different genera was also detected in fossil forms of Spatangus, Propespatagus nov. gen., and Sardospatangus (†) from the European Oligo-Miocene sedimentary rocks of Germany; the Miocene of Ukraine, Italy, and North Africa; the Plio-Pleistocene of Italy; and the Mio-Pliocene of Florida (USA). The new data can help in addressing taxonomic ambiguities within echinoids, as well as in improving species identification, and hence biodiversity assessments in the Mediterranean region.

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