Project 3497: J. C. Lamsdell. 2020. The phylogeny and systematics of Xiphosura. PeerJ. (In Press)
Specimen: † Lunataspis aurora (unvouchered)
View: Dorsal

Abstract

Xiphosurans are aquatic chelicerates with a fossil record extending into the Early Ordovician and known from a total of 88 described species, four of which are extant. Known for their apparent morphological conservatism, for which they have gained notoriety as supposed ‘living fossils’, recent analyses have demonstrated xiphosurans to have an ecologically diverse evolutionary history, with several groups moving into non-marine environments and developing morphologies markedly different from those of the modern species. The combination of their long evolutionary and complex ecological history along with their paradoxical patterns of morphological stasis in some clades and experimentation among others has resulted in Xiphosura being of particular interest for macroevolutionary study. Phylogenetic analyses have shown the current taxonomic framework for Xiphosura – set out in the Treatise of Invertebrate Paleontology in 1955 – to be outdated and in need of revision, with several common genera such as Paleolimulus Dunbar, 1923 and Limulitella Størmer, 1952 acting as wastebasket taxon. Here, an expanded xiphosuran phylogeny is presented, comprising 58 xiphosuran species as part of a 158 taxon chelicerate matrix coded for 259 characters. Analysing the matrix under both Bayesian inference and parsimony optimisation criteria retrieves a concordant tree topology that forms the basis of a genus-level systematic revision of xiphosuran taxonomy. The genera Euproops Meek, 1867 , Belinurus König, 1820, Paleolimulus, Limulitella, and Limulus are demonstrated to be non-monophyletic and the previously synonymized genera Koenigiella Raymond, 1944 and Prestwichianella Cockerell, 1905 are shown to be valid. In addition, nine new genera (Andersoniella gen. nov., Macrobelinurus gen. nov., and Parabelinurus gen. nov. in Belinurina; Norilimulus gen. nov. in Paleolimulidae; Batracholimulus gen. nov. and Boeotiaspis gen. nov. in Austrolimulidae; and Allolimulus gen. nov., Keuperlimulus gen. nov., and Volanalimulus gen. nov. in Limulidae) are erected to accommodate xiphosuran species not encompassed by existing genera. One new species, Volanalimulus madagascarensis gen. et sp. nov., is also described. Three putative xiphosuran genera – Elleria Raymond, 1944, Archeolimulus Chlupáč, 1963, and Drabovaspis Chlupáč, 1963 – are determined to be non-xiphosuran arthropods and as such are removed from Xiphosura. The priority of Belinurus König, 1820 over Bellinurus Pictet, 1846 is also confirmed. This work is critical for facilitating the study of the xiphosuran fossil record and is the first step in resolving longstanding questions regarding the geographic distribution of the modern horseshoe crab species and whether they truly represent ‘living fossils’. Understanding the long evolutionary history of Xiphosura is vital for interpreting how the modern species may respond to environmental change and in guiding conservation efforts.


Project DOI: 10.7934/P3497, http://dx.doi.org/10.7934/P3497
This project containsMatrices
  • 1 Media
  • 1 Matrix
  • 160 Taxa
  • 2 Specimens
  • 259 Characters
Total size of project's media files: 14.48M

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Total scored cells: 27773
Total media associated with cells: 0
Total labels associated with cell media: 0
Characters
Total characters: 259
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 620
Total character states with associated media: 0
Total character states with media with labels:0
Total unordered/ordered characters:259/0
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MorphoBank Project 3497

    This research
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    Authors' Institutions

    • West Virginia University



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    James Lamsdell
    Project Administrator
    160212590027773
    (21419, 0, 6354)
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    Taxonomic Overview for Matrix 'M26157' (160 Taxa)

    taxon unscored
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    [1] † Fuxianhuia protensa
    Taxon name last Modified on 07/02/14
    35149149075001
    [2] † Leanchoilia illecebrosa
    Taxon name last Modified on 07/02/14
    36163163060001
    [3] † Alalcomenaeus cambricus
    Taxon name last Modified on 07/02/14
    28170170061001
    [4] † Yohoia tenuis
    Taxon name last Modified on 07/02/14
    70133133056001
    [5] † Emeraldella brocki
    Taxon name last Modified on 07/02/14
    21165165073001
    [6] † Sydneyia inexpectans
    Taxon name last Modified on 07/02/14
    42147147070001
    [7] † Olenoides serratus
    Taxon name last Modified on 07/02/14
    13188188058001
    [8] Pycnogonum litorale
    Taxon name last Modified on 07/02/14
    81021020149001
    [9] † Haliestes dasos
    Taxon name last Modified on 07/02/14
    3985850135001
    [10] † Palaeoisopus problematicus
    Taxon name last Modified on 07/02/14
    221161160121001
    [11] † Offacolus kingi
    Taxon name last Modified on 07/02/14
    87131131041001
    [12] † Dibasterium durgae
    Taxon name last Modified on 07/02/14
    57149149053001
    [13] † Anderella parva
    Taxon name last Modified on 08/20/14
    1568787016001
    [14] † Weinbergina opitzi
    Taxon name last Modified on 07/02/14
    68161161030001
    [15] † Camanchia grovensis
    Taxon name last Modified on 07/02/14
    1479595017001
    [16] † Venustulus waukeshaensis
    Taxon name last Modified on 07/02/14
    128111111020001
    [17] † Legrandella lombardi
    Taxon name last Modified on 07/02/14
    137107107015001
    [18] † Bembicosoma pomphicus
    Taxon name last Modified on 07/02/14
    1598585015001
    [19] † Bunodes lunula
    Taxon name last Modified on 11/13/20
    1568686017001
    [20] † Limuloides limuloides
    Taxon name last Modified on 07/02/14
    1489797014001
    [21] † Pasternakevia podolica
    Taxon name last Modified on 07/02/14
    1578787015001
    [22] † Cyamocephalus loganensis
    Taxon name last Modified on 07/02/14
    1509494015001
    [23] † Pseudoniscus roosevelti
    Taxon name last Modified on 07/02/14
    1568888015001
    [24] † Winneshiekia youngae
    Taxon name last Modified on 07/16/18
    1568989014001
    [25] † Houia yueya
    Taxon name last Modified on 05/25/15
    1448989026001
    [26] † Euproops anthrax
    Taxon name last Modified on 07/02/14
    98138138023001
    [27] † Euproops mariae
    Taxon name last Modified on 07/17/14
    1688080011001
    [28] † Euproops danae
    Taxon name last Modified on 07/02/14
    93143143023001
    [29] † Euproops rotundatus
    Taxon name last Modified on 07/17/14
    146101101012001
    [30] † Euproops sp
    Taxon name last Modified on 07/02/14
    91141141027001
    [31] † Bellinurus lunatus
    Taxon name last Modified on 07/18/14
    1489898013001
    [32] † Bellinurus arcuatus
    Taxon name last Modified on 07/18/14
    1499797013001
    [33] † Bellinurus reginae
    Taxon name last Modified on 07/18/14
    1499696014001
    [34] † Bellinurus truemanii
    Taxon name last Modified on 07/18/14
    1509595014001
    [35] † Bellinurus bellulus
    Taxon name last Modified on 07/18/14
    1539292014001
    [36] † Bellinurus trilobitoides
    Taxon name last Modified on 07/02/14
    1499696014001
    [37] † Anacontium carpenteri
    Taxon name last Modified on 07/16/14
    220373702001
    [38] † Liomesaspis laevis
    Taxon name last Modified on 07/16/14
    1519090018001
    [39] † Stilpnocephalus pontebbanus
    Taxon name last Modified on 07/27/20
    2222727010001
    [40] † Alanops magnifica
    Taxon name last Modified on 07/16/14
    59157157043001
    [41] † Pringlia birtwelli
    Taxon name last Modified on 07/17/14
    1627575022001
    [42] † Kasibelinurus amoricum
    Taxon name last Modified on 07/02/14
    1549393012001
    [43] † Kasibelinurus randalli
    Taxon name last Modified on 07/02/14
    1648383012001
    [44] † Pickettia carteri
    Taxon name last Modified on 07/27/20
    1628484013001
    [45] † Lunataspis aurora
    Taxon name last Modified on 07/02/14
    1429999018001
    [46] Limulus polyphemus
    Taxon name last Modified on 07/02/14
    0219219040001
    [47] † Limulus darwini
    Taxon name last Modified on 07/27/20
    1628080017001
    [48] † Limulus coffini
    Taxon name last Modified on 07/14/14
    1994646014001
    [49] † Crenatolimulus paluxyensis
    Taxon name last Modified on 07/14/14
    1598484016001
    [50] Carcinoscorpius rotundicauda
    Taxon name last Modified on 07/03/14
    0220220039001
    [51] † Heterolimulus gadeai
    Taxon name last Modified on 07/16/18
    1499494016001
    [52] Tachypleus gigas
    Taxon name last Modified on 07/03/14
    0220220039001
    [53] † Tachypleus decheni
    Taxon name last Modified on 07/16/18
    1489595016001
    [54] † Tachypleus syriacus
    Taxon name last Modified on 07/16/18
    143101100016001
    [55] Tachypleus tridentatus
    Taxon name last Modified on 07/02/14
    0220220039001
    [56] † Yunnanolimulus luopingensis
    Taxon name last Modified on 07/11/14
    30189189040001
    [57] † Xaniopyramis linseyi
    Taxon name last Modified on 07/10/14
    187676705001
    [58] † Moravurus rehori
    Taxon name last Modified on 07/27/20
    231232305001
    [59] † Paleolimulus woodae
    Taxon name last Modified on 07/11/18
    1578989013001
    [60] † Paleolimulus signatus
    Taxon name last Modified on 07/02/14
    46183183030001
    [61] † Paleolimulus kunguricus
    Taxon name last Modified on 07/16/18
    141104104014001
    [62] † Bellinuroopsis rossicus
    Taxon name last Modified on 07/18/14
    1529393014001
    [63] † Rolfeia fouldenensis
    Taxon name last Modified on 07/10/14
    143102102014001
    [64] † Tarracolimulus rieki
    Taxon name last Modified on 07/14/14
    1558989015001
    [65] † Casterolimulus kletti
    Taxon name last Modified on 07/16/14
    221363602001
    [66] † Limulitella vicensis
    Taxon name last Modified on 07/21/14
    1757272012001
    [67] † Victalimulus mcqueeni
    Taxon name last Modified on 07/16/14
    116125125018001
    [68] † Limulus woodwardi
    Taxon name last Modified on 07/13/18
    222353502001
    [69] † Mesolimulus crespelli
    Taxon name last Modified on 07/15/14
    1647979016001
    [70] † Mesolimulus tafraoutensis
    Taxon name last Modified on 07/27/20
    1558888016001
    [71] † Mesolimulus walchi
    Taxon name last Modified on 07/02/14
    57173173029001
    [72] † Mesolimulus sibiricus
    Taxon name last Modified on 07/27/20
    1618383015001
    [73] † Panduralimulus babcocki
    Taxon name last Modified on 07/11/14
    1429898019001
    [74] † Tasmaniolimulus patersoni
    Taxon name last Modified on 07/27/20
    1608080019001
    [75] † Austrolimulus fletcheri
    Taxon name last Modified on 07/11/14
    1499191019001
    [76] † Vaderlimulus tricki
    Taxon name last Modified on 07/11/18
    1479292020001
    [77] † Psammolimulus gottingensis
    Taxon name last Modified on 07/11/14
    113122122024001
    [78] † Limulitella bronni
    Taxon name last Modified on 07/17/14
    1499090020001
    [79] † Limulitella tejraensis
    Taxon name last Modified on 07/11/18
    1607979020001
    [80] † Limulitella henkeli
    Taxon name last Modified on 07/17/14
    1697575015001
    [81] † Madagascan limulid
    Taxon name last Modified on 07/23/19
    1657979015001
    [82] † Paleolimulus fuchsbergensis
    Taxon name last Modified on 07/11/14
    1568484019001
    [83] † Dubbolimulus peetae
    Taxon name last Modified on 07/16/18
    1469393020001
    [84] † Paleolimulus longispinus
    Taxon name last Modified on 07/15/14
    137100100022001
    [85] † Paleolimulus jakovlevi
    Taxon name last Modified on 07/27/20
    1657878016001
    [86] † Valloisella lievinensis
    Taxon name last Modified on 07/16/14
    1469797016001
    [87] † Stoermeropterus conicus
    Taxon name last Modified on 07/02/14
    25199199035001
    [88] † Moselopterus ancylotelson
    Taxon name last Modified on 07/02/14
    24195195040001
    [89] † Onychopterella kokomoensis
    Taxon name last Modified on 07/02/14
    33190190036001
    [90] † Eurypterus tetragonophthalmus
    Taxon name last Modified on 07/02/14
    10214214035001
    [91] † Erieopterus macrophthalmus
    Taxon name last Modified on 07/02/14
    33191191035001
    [92] † Strobilopterus proteus
    Taxon name last Modified on 07/02/14
    47188188024001
    [93] † Dolichopterus macrocheirus
    Taxon name last Modified on 07/02/14
    23202202034001
    [94] † Carcinosoma newlini
    Taxon name last Modified on 07/02/14
    31189189039001
    [95] † Megalograptus ohioensis
    Taxon name last Modified on 07/02/14
    22202202035001
    [96] † Mixopterus kiaeri
    Taxon name last Modified on 07/02/14
    32188188039001
    [97] † Orcanopterus manitoulinensis
    Taxon name last Modified on 07/02/14
    73146146040001
    [98] † Adelophthalmus sievertsi
    Taxon name last Modified on 07/02/14
    46176176037001
    [99] † Nanahughmilleria norvegica
    Taxon name last Modified on 07/02/14
    24199199036001
    [100] † Hughmilleria socialis
    Taxon name last Modified on 07/02/14
    23196196040001
    [101] † Slimonia acuminata
    Taxon name last Modified on 07/02/14
    25194194040001
    [102] † Erettopterus bilobus
    Taxon name last Modified on 07/02/14
    25192192042001
    [103] † Pterygotus anglicus
    Taxon name last Modified on 07/02/14
    31185185043001
    [104] † Brachyopterus stubblefieldi
    Taxon name last Modified on 07/02/14
    71151151037001
    [105] † Rhenopterus diensti
    Taxon name last Modified on 07/02/14
    24199199036001
    [106] † Parastylonurus ornatus
    Taxon name last Modified on 07/02/14
    34189189036001
    [107] † Laurieipterus elegans
    Taxon name last Modified on 07/02/14
    94129129036001
    [108] † Kokomopterus longicaudatus
    Taxon name last Modified on 07/02/14
    44178178037001
    [109] † Hardieopterus macrophthalmus
    Taxon name last Modified on 07/02/14
    74151151034001
    [110] † Drepanopterus pentlandicus
    Taxon name last Modified on 07/02/14
    28199198033001
    [111] † Hibbertopterus scouleri
    Taxon name last Modified on 07/02/14
    31196196032001
    [112] † Megarachne servinei
    Taxon name last Modified on 07/02/14
    1736362024001
    [113] † Chasmataspis laurencii
    Taxon name last Modified on 07/02/14
    112122122025001
    [114] † Loganamaraspis dunlopi
    Taxon name last Modified on 07/02/14
    97129129033001
    [115] † Achanarraspis reedi
    Taxon name last Modified on 07/02/14
    1538686020001
    [116] † Heteroaspis stoermeri
    Taxon name last Modified on 07/02/14
    1449191024001
    [117] † Diploaspis casteri
    Taxon name last Modified on 07/02/14
    107125125027001
    [118] † Diploaspis muelleri
    Taxon name last Modified on 07/02/14
    1488787024001
    [119] † Dvulikiaspis menneri
    Taxon name last Modified on 07/02/14
    122106106031001
    [120] † Octoberaspis ushakovi
    Taxon name last Modified on 07/02/14
    65160160034001
    [121] Centruroides vittatus
    Taxon name last Modified on 11/13/20
    1198198060001
    [122] Androctonus australis
    Taxon name last Modified on 07/02/14
    0198198061001
    [123] † Proscorpius osborni
    Taxon name last Modified on 07/02/14
    29169169061001
    [124] † Palaeophonus nuncius
    Taxon name last Modified on 07/02/14
    54146146059001
    [125] Caddo agilis
    Taxon name last Modified on 11/13/20
    7177177075001
    [126] Chileogovea oedipus
    Taxon name last Modified on 11/13/20
    15173173071001
    [127] Cyphophthalmus duricorius
    Taxon name last Modified on 07/02/14
    15173173071001
    [128] Gonyleptes horridus
    Taxon name last Modified on 07/02/14
    7178178074001
    [129] Leiobunum rotundum
    Taxon name last Modified on 07/02/14
    7177177075001
    [130] Sclerobunus robustus
    Taxon name last Modified on 07/02/14
    7178178074001
    [131] Allothrombium simoni
    Taxon name last Modified on 07/02/14
    16160159084001
    [132] Amblyomma americanum
    Taxon name last Modified on 11/13/20
    14158157088001
    [133] Neocarus texanus
    Taxon name last Modified on 07/02/14
    8167167084001
    [134] Cryptocellus goodnighti
    Taxon name last Modified on 07/02/14
    4180180075001
    [135] Poliochera punctulata
    Taxon name last Modified on 07/02/14
    59126126074001
    [136] Ricinoides atewa
    Taxon name last Modified on 07/02/14
    4180180075001
    [137] † Anthracomartus hindi
    Taxon name last Modified on 07/02/14
    31157157071001
    [138] † Eophrynus prestvicii
    Taxon name last Modified on 07/02/14
    32159159068001
    [139] † Gilboarachne griersoni
    Taxon name last Modified on 07/02/14
    20167167072001
    [140] † Palaeocharinus sp.
    Taxon name last Modified on 07/02/14
    8179179072001
    [141] † Mesotarbus peteri
    Taxon name last Modified on 07/02/14
    51136136072001
    [142] † Plesiosiro madeleyi
    Taxon name last Modified on 07/02/14
    43145145071001
    [143] Eukoenenia mirabilis
    Taxon name last Modified on 07/02/14
    3181180076001
    [144] Prokoenenia wheeleri
    Taxon name last Modified on 07/02/14
    3181180076001
    [145] Mastigoproctus giganteus
    Taxon name last Modified on 11/13/20
    3190190066001
    [146] Thelyphonus caudatus
    Taxon name last Modified on 07/02/14
    3186186070001
    [147] Hubbardia pentapeltis
    Taxon name last Modified on 07/02/14
    3186185071001
    [148] Schizomus crassicaudatus
    Taxon name last Modified on 07/02/14
    3186185071001
    [149] † Permarachne novokshonovi
    Taxon name last Modified on 07/02/14
    119101101039001
    [150] † Attercopus filmbriunguis
    Taxon name last Modified on 07/02/14
    1736868018001
    [151] Liphistius malayanus
    Taxon name last Modified on 07/02/14
    3175175081001
    [152] Aphonopelma hentzi
    Taxon name last Modified on 11/13/20
    3166166090001
    [153] Hypochilus pococki
    Taxon name last Modified on 11/13/20
    3166166090001
    [154] Cupienius getazi
    Taxon name last Modified on 07/02/14
    4165165090001
    [155] Neobisium maritimum
    Taxon name last Modified on 07/02/14
    10166165084001
    [156] Chelifer cancroides
    Taxon name last Modified on 11/13/20
    10165164085001
    [157] Eremocosta striata
    Taxon name last Modified on 07/02/14
    10162161088001
    [158] Galeodes armeniacus
    Taxon name last Modified on 07/02/14
    12161160087001
    [159] Phrynus marginemaculatus
    Taxon name last Modified on 07/02/14
    4182182073001
    [160] Charinus victori
    Taxon name last Modified on 07/02/14
    4181181074001


    Project downloads

    type number of downloads Individual items downloaded (where applicable)
    Total downloads from project175
    Project downloads115
    Matrix downloads59Xiph focus (59 downloads);
    Media downloads1M736137 (1 download);