A new limulid genus from the Strelovec Formation (Middle Triassic, Anisian) of northern Slovenia

Horseshoe crabs are an archetypal chelicerate group with a fossil record extending back to Early Ordovician time. Although extensively studied, the group generally has a low diversity across the Phanerozoic Eonothem. Here, we expand the known diversity of true horseshoe crabs (Xiphosurida) by the description of a new taxon from the Middle Triassic Strelovec Formation of the Slovenian Alps. The mostly complete fossil is preserved as an external mould and assigned to the family Limulidae Zittel, 1881 as Sloveniolimulus rudkini, n. gen., n. sp. The use of landmark and semilandmark geometric morphometrics is explored to corroborate the systematic palaeontology and suggests that the new genus and species are valid. We also provide the first quantitative evidence for the extensive diversity of Triassic horseshoe crabs. We suggest that Triassic horseshoe crabs likely filled many ecological niches left vacant after the end-Permian extinction.

Phylogeny of Paleozoic limbed vertebrates reassessed through revision and expansion of the largest published relevant data matrix

The largest published phylogenetic analysis of early limbed vertebrates (Ruta M, Coates MI. 2007.Journal of Systematic Palaeontology5:69–122) recovered, for example, Seymouriamorpha, Diadectomorpha and (in some trees) Caudata as paraphyletic and found the “temnospondyl hypothesis” on the origin of Lissamphibia (TH) to be more parsimonious than the “lepospondyl hypothesis” (LH)—though only, as we show, by one step. We report 4,200 misscored cells, over half of them due to typographic and similar accidental errors. Further, some characters were duplicated; some had only one described state; for one, most taxa were scored after presumed relatives. Even potentially continuous characters were unordered, the effects of ontogeny were not sufficiently taken into account, and data published after 2001 were mostly excluded. After these issues are improved—we document and justify all changes to the matrix—but no characters are added, we find (Analysis R1) much longer trees with, for example, monophyletic Caudata, Diadectomorpha and (in some trees) Seymouriamorpha;Ichthyostegaeither crownward or rootward ofAcanthostega; and Anthracosauria either crownward or rootward of Temnospondyli. The LH is nine steps shorter than the TH (R2; constrained) and 12 steps shorter than the “polyphyly hypothesis” (PH—R3; constrained).Brachydectes(Lysorophia) is not found next to Lissamphibia; instead, a large clade that includes the adelogyrinids, urocordylid “nectrideans” and aïstopods occupies that position. As expected from the taxon/character ratio, most bootstrap values are low. Adding 56 terminal taxa to the original 102 increases the resolution (and decreases most bootstrap values). The added taxa range in completeness from complete articulated skeletons to an incomplete lower jaw. Even though the lissamphibian-like temnospondylsGerobatrachus,MicropholisandTungussogyrinusand the extremely peramorphic salamanderChelotritonare added, the difference between LH (R4; unconstrained) and TH (R5) rises to 10 steps, that between LH and PH (R6) to 15; the TH also requires several more regains of lost bones than the LH.Casineria, in which we tentatively identify a postbranchial lamina, emerges rather far from amniote origins in a gephyrostegid-chroniosuchian grade. Bayesian inference (Analysis EB, settings as in R4) mostly agrees with R4. High posterior probabilities are found for Lissamphibia (1.00) and the LH (0.92); however, many branches remain weakly supported, and most are short, as expected from the small character sample. We discuss phylogeny, approaches to coding, methods of phylogenetics (Bayesian inference vs. equally weighted vs. reweighted parsimony), some character complexes (e.g. preaxial/postaxial polarity in limb development), and prospects for further improvement of this matrix. Even in its revised state, the matrix cannot provide a robust assessment of the phylogeny of early limbed vertebrates. Sufficient improvement will be laborious—but not difficult.