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the current taxonomic framework for Xiphosura – set out in the Treatise of Invertebrate Paleontology in 1955 – to be outdated and in need of revision, with several common genera such as Paleolimulus Dunbar, 1923 and Limulitella Størmer, 1952 acting as wastebasket taxon.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

Download Project SDD File
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
    supported by

    Authors' Institutions

    • West Virginia University



    Members

    member name taxa specimens media chars character
    media
    labels
    cell scorings
    (scored, NPA, "-")
    cell
    medialabels
    rules
    James Lamsdell
    Project Administrator
    Last logged in 11/13/20
    160212590027773
    (21419, 0, 6354)
    000


    Taxonomic Overview for Matrix 'M26157' (160 Taxa)

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


    Project views

    type number of views Individual items viewed (where applicable)
    Total project views179
    Project overview31
    Matrix views15Matrix landing page (10 views); Xiph focus (5 views);
    Media views42Media search (36 views); M736137 (6 views);
    Views for media list8
    Bibliography5
    Specimen list22
    Taxon list56




    Project downloads

    type number of downloads Individual items downloaded (where applicable)
    Total downloads from project4
    Project downloads3
    Matrix downloads1Xiph focus (1 download);