[Palaeontology, Vol. 59, Part 2, 2016, pp. 181–194] HORSESHOE CRAB PHYLOGENY AND INDEPENDENT COLONIZATIONS OF FRESH WATER: ECOLOGICAL INVASION AS A DRIVER FOR MORPHOLOGICAL INNOVATION by JAMES C. LAMSDELL Department of Geology and Geophysics, Yale University, PO Box 208109, New Haven, CT 06520-8109, USA; e-mail:
[email protected] Typescript received 10 August 2015; accepted in revised form 8 October 2015 Abstract: Xiphosurids are an archaic group of aquatic the group throughout the Phanerozoic. Xiphosurids are chelicerate arthropods, generally known by the colloquial shown to have invaded non-marine environments indepen- misnomer of ‘horseshoe crabs’. Known from marine envi- dently at least five times throughout their evolutionary ronments as far back as the early Ordovician, horseshoe history, twice resulting in the radiation of major clades – crabs are generally considered ‘living fossils’ – descendants bellinurines and austrolimulids – that occupied novel of a bradytelic lineage exhibiting little morphological or regions of morphospace. These clades show a convergent ecological variation throughout geological time. However, ecological pattern of differentiation, speciation and subse- xiphosurids are known from freshwater sediments in the quent extinction. Horseshoe crabs are shown to have a Palaeozoic and Mesozoic; furthermore, the contention that more dynamic and complex evolutionary history than previ- xiphosurids show little morphological variation has never ously supposed, with the extant species representing only a been tested empirically. Attempts to test this are hampered fraction of the group’s past ecological and morphological by the lack of a modern phylogenetic framework with diversity. which to explore different evolutionary scenarios. Here, I present a phylogenetic analysis of Xiphosurida and explore Key words: bradytely, convergence, ecology, morphospace, patterns of morphospace and environmental occupation of niche conservatism, Xiphosurida.