Morphobank

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the current taxonomic framework for Xiphosura – set out in the Treatise of Invertebrate Paleontology in 1955 – to be outdated and in need of revision, with several common genera such as Paleolimulus Dunbar, 1923 and Limulitella Størmer, 1952 acting as wastebasket taxon.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

Currently Viewing:
MorphoBank Project 3497

Creation Date:
11 July 2019
Publication Date:
13 November 2020
Media downloads: 1
Matrix downloads: 54

This research
supported by

Authors' Institutions

West Virginia University

Members

James Lamsdell
Project Administrator

160

259

27773
(21419, 0, 6354)

Taxonomic Overview for Matrix 'M26157' (160 Taxa)

taxon	unscored cells	scored cells	no cell support	NPA cells	"-" cells	cell images	labels on cell images	member access
[1] † Fuxianhuia protensa Taxon name last Modified on 07/02/14	35	149	149	0	75	0	0	1
[2] † Leanchoilia illecebrosa Taxon name last Modified on 07/02/14	36	163	163	0	60	0	0	1
[3] † Alalcomenaeus cambricus Taxon name last Modified on 07/02/14	28	170	170	0	61	0	0	1
[4] † Yohoia tenuis Taxon name last Modified on 07/02/14	70	133	133	0	56	0	0	1
[5] † Emeraldella brocki Taxon name last Modified on 07/02/14	21	165	165	0	73	0	0	1
[6] † Sydneyia inexpectans Taxon name last Modified on 07/02/14	42	147	147	0	70	0	0	1
[7] † Olenoides serratus Taxon name last Modified on 07/02/14	13	188	188	0	58	0	0	1
[8] Pycnogonum litorale Taxon name last Modified on 07/02/14	8	102	102	0	149	0	0	1
[9] † Haliestes dasos Taxon name last Modified on 07/02/14	39	85	85	0	135	0	0	1
[10] † Palaeoisopus problematicus Taxon name last Modified on 07/02/14	22	116	116	0	121	0	0	1
[11] † Offacolus kingi Taxon name last Modified on 07/02/14	87	131	131	0	41	0	0	1
[12] † Dibasterium durgae Taxon name last Modified on 07/02/14	57	149	149	0	53	0	0	1
[13] † Anderella parva Taxon name last Modified on 08/20/14	156	87	87	0	16	0	0	1
[14] † Weinbergina opitzi Taxon name last Modified on 07/02/14	68	161	161	0	30	0	0	1
[15] † Camanchia grovensis Taxon name last Modified on 07/02/14	147	95	95	0	17	0	0	1
[16] † Venustulus waukeshaensis Taxon name last Modified on 07/02/14	128	111	111	0	20	0	0	1
[17] † Legrandella lombardi Taxon name last Modified on 07/02/14	137	107	107	0	15	0	0	1
[18] † Bembicosoma pomphicus Taxon name last Modified on 07/02/14	159	85	85	0	15	0	0	1
[19] † Bunodes lunula Taxon name last Modified on 11/13/20	156	86	86	0	17	0	0	1
[20] † Limuloides limuloides Taxon name last Modified on 07/02/14	148	97	97	0	14	0	0	1
[21] † Pasternakevia podolica Taxon name last Modified on 07/02/14	157	87	87	0	15	0	0	1
[22] † Cyamocephalus loganensis Taxon name last Modified on 07/02/14	150	94	94	0	15	0	0	1
[23] † Pseudoniscus roosevelti Taxon name last Modified on 07/02/14	156	88	88	0	15	0	0	1
[24] † Winneshiekia youngae Taxon name last Modified on 07/16/18	156	89	89	0	14	0	0	1
[25] † Houia yueya Taxon name last Modified on 05/25/15	144	89	89	0	26	0	0	1
[26] † Euproops anthrax Taxon name last Modified on 07/02/14	98	138	138	0	23	0	0	1
[27] † Euproops mariae Taxon name last Modified on 07/17/14	168	80	80	0	11	0	0	1
[28] † Euproops danae Taxon name last Modified on 07/02/14	93	143	143	0	23	0	0	1
[29] † Euproops rotundatus Taxon name last Modified on 07/17/14	146	101	101	0	12	0	0	1
[30] † Euproops sp Taxon name last Modified on 07/02/14	91	141	141	0	27	0	0	1
[31] † Bellinurus lunatus Taxon name last Modified on 07/18/14	148	98	98	0	13	0	0	1
[32] † Bellinurus arcuatus Taxon name last Modified on 07/18/14	149	97	97	0	13	0	0	1
[33] † Bellinurus reginae Taxon name last Modified on 07/18/14	149	96	96	0	14	0	0	1
[34] † Bellinurus truemanii Taxon name last Modified on 07/18/14	150	95	95	0	14	0	0	1
[35] † Bellinurus bellulus Taxon name last Modified on 07/18/14	153	92	92	0	14	0	0	1
[36] † Bellinurus trilobitoides Taxon name last Modified on 07/02/14	149	96	96	0	14	0	0	1
[37] † Anacontium carpenteri Taxon name last Modified on 07/16/14	220	37	37	0	2	0	0	1
[38] † Liomesaspis laevis Taxon name last Modified on 07/16/14	151	90	90	0	18	0	0	1
[39] † Stilpnocephalus pontebbanus Taxon name last Modified on 07/27/20	222	27	27	0	10	0	0	1
[40] † Alanops magnifica Taxon name last Modified on 07/16/14	59	157	157	0	43	0	0	1
[41] † Pringlia birtwelli Taxon name last Modified on 07/17/14	162	75	75	0	22	0	0	1
[42] † Kasibelinurus amoricum Taxon name last Modified on 07/02/14	154	93	93	0	12	0	0	1
[43] † Kasibelinurus randalli Taxon name last Modified on 07/02/14	164	83	83	0	12	0	0	1
[44] † Pickettia carteri Taxon name last Modified on 07/27/20	162	84	84	0	13	0	0	1
[45] † Lunataspis aurora Taxon name last Modified on 07/02/14	142	99	99	0	18	0	0	1
[46] Limulus polyphemus Taxon name last Modified on 07/02/14	0	219	219	0	40	0	0	1
[47] † Limulus darwini Taxon name last Modified on 07/27/20	162	80	80	0	17	0	0	1
[48] † Limulus coffini Taxon name last Modified on 07/14/14	199	46	46	0	14	0	0	1
[49] † Crenatolimulus paluxyensis Taxon name last Modified on 07/14/14	159	84	84	0	16	0	0	1
[50] Carcinoscorpius rotundicauda Taxon name last Modified on 07/03/14	0	220	220	0	39	0	0	1
[51] † Heterolimulus gadeai Taxon name last Modified on 07/16/18	149	94	94	0	16	0	0	1
[52] Tachypleus gigas Taxon name last Modified on 07/03/14	0	220	220	0	39	0	0	1
[53] † Tachypleus decheni Taxon name last Modified on 07/16/18	148	95	95	0	16	0	0	1
[54] † Tachypleus syriacus Taxon name last Modified on 07/16/18	143	101	100	0	16	0	0	1
[55] Tachypleus tridentatus Taxon name last Modified on 07/02/14	0	220	220	0	39	0	0	1
[56] † Yunnanolimulus luopingensis Taxon name last Modified on 07/11/14	30	189	189	0	40	0	0	1
[57] † Xaniopyramis linseyi Taxon name last Modified on 07/10/14	187	67	67	0	5	0	0	1
[58] † Moravurus rehori Taxon name last Modified on 07/27/20	231	23	23	0	5	0	0	1
[59] † Paleolimulus woodae Taxon name last Modified on 07/11/18	157	89	89	0	13	0	0	1
[60] † Paleolimulus signatus Taxon name last Modified on 07/02/14	46	183	183	0	30	0	0	1
[61] † Paleolimulus kunguricus Taxon name last Modified on 07/16/18	141	104	104	0	14	0	0	1
[62] † Bellinuroopsis rossicus Taxon name last Modified on 07/18/14	152	93	93	0	14	0	0	1
[63] † Rolfeia fouldenensis Taxon name last Modified on 07/10/14	143	102	102	0	14	0	0	1
[64] † Tarracolimulus rieki Taxon name last Modified on 07/14/14	155	89	89	0	15	0	0	1
[65] † Casterolimulus kletti Taxon name last Modified on 07/16/14	221	36	36	0	2	0	0	1
[66] † Limulitella vicensis Taxon name last Modified on 07/21/14	175	72	72	0	12	0	0	1
[67] † Victalimulus mcqueeni Taxon name last Modified on 07/16/14	116	125	125	0	18	0	0	1
[68] † Limulus woodwardi Taxon name last Modified on 07/13/18	222	35	35	0	2	0	0	1
[69] † Mesolimulus crespelli Taxon name last Modified on 07/15/14	164	79	79	0	16	0	0	1
[70] † Mesolimulus tafraoutensis Taxon name last Modified on 07/27/20	155	88	88	0	16	0	0	1
[71] † Mesolimulus walchi Taxon name last Modified on 07/02/14	57	173	173	0	29	0	0	1
[72] † Mesolimulus sibiricus Taxon name last Modified on 07/27/20	161	83	83	0	15	0	0	1
[73] † Panduralimulus babcocki Taxon name last Modified on 07/11/14	142	98	98	0	19	0	0	1
[74] † Tasmaniolimulus patersoni Taxon name last Modified on 07/27/20	160	80	80	0	19	0	0	1
[75] † Austrolimulus fletcheri Taxon name last Modified on 07/11/14	149	91	91	0	19	0	0	1
[76] † Vaderlimulus tricki Taxon name last Modified on 07/11/18	147	92	92	0	20	0	0	1
[77] † Psammolimulus gottingensis Taxon name last Modified on 07/11/14	113	122	122	0	24	0	0	1
[78] † Limulitella bronni Taxon name last Modified on 07/17/14	149	90	90	0	20	0	0	1
[79] † Limulitella tejraensis Taxon name last Modified on 07/11/18	160	79	79	0	20	0	0	1
[80] † Limulitella henkeli Taxon name last Modified on 07/17/14	169	75	75	0	15	0	0	1
[81] † Madagascan limulid Taxon name last Modified on 07/23/19	165	79	79	0	15	0	0	1
[82] † Paleolimulus fuchsbergensis Taxon name last Modified on 07/11/14	156	84	84	0	19	0	0	1
[83] † Dubbolimulus peetae Taxon name last Modified on 07/16/18	146	93	93	0	20	0	0	1
[84] † Paleolimulus longispinus Taxon name last Modified on 07/15/14	137	100	100	0	22	0	0	1
[85] † Paleolimulus jakovlevi Taxon name last Modified on 07/27/20	165	78	78	0	16	0	0	1
[86] † Valloisella lievinensis Taxon name last Modified on 07/16/14	146	97	97	0	16	0	0	1
[87] † Stoermeropterus conicus Taxon name last Modified on 07/02/14	25	199	199	0	35	0	0	1
[88] † Moselopterus ancylotelson Taxon name last Modified on 07/02/14	24	195	195	0	40	0	0	1
[89] † Onychopterella kokomoensis Taxon name last Modified on 07/02/14	33	190	190	0	36	0	0	1
[90] † Eurypterus tetragonophthalmus Taxon name last Modified on 07/02/14	10	214	214	0	35	0	0	1
[91] † Erieopterus macrophthalmus Taxon name last Modified on 07/02/14	33	191	191	0	35	0	0	1
[92] † Strobilopterus proteus Taxon name last Modified on 07/02/14	47	188	188	0	24	0	0	1
[93] † Dolichopterus macrocheirus Taxon name last Modified on 07/02/14	23	202	202	0	34	0	0	1
[94] † Carcinosoma newlini Taxon name last Modified on 07/02/14	31	189	189	0	39	0	0	1
[95] † Megalograptus ohioensis Taxon name last Modified on 07/02/14	22	202	202	0	35	0	0	1
[96] † Mixopterus kiaeri Taxon name last Modified on 07/02/14	32	188	188	0	39	0	0	1
[97] † Orcanopterus manitoulinensis Taxon name last Modified on 07/02/14	73	146	146	0	40	0	0	1
[98] † Adelophthalmus sievertsi Taxon name last Modified on 07/02/14	46	176	176	0	37	0	0	1
[99] † Nanahughmilleria norvegica Taxon name last Modified on 07/02/14	24	199	199	0	36	0	0	1
[100] † Hughmilleria socialis Taxon name last Modified on 07/02/14	23	196	196	0	40	0	0	1
[101] † Slimonia acuminata Taxon name last Modified on 07/02/14	25	194	194	0	40	0	0	1
[102] † Erettopterus bilobus Taxon name last Modified on 07/02/14	25	192	192	0	42	0	0	1
[103] † Pterygotus anglicus Taxon name last Modified on 07/02/14	31	185	185	0	43	0	0	1
[104] † Brachyopterus stubblefieldi Taxon name last Modified on 07/02/14	71	151	151	0	37	0	0	1
[105] † Rhenopterus diensti Taxon name last Modified on 07/02/14	24	199	199	0	36	0	0	1
[106] † Parastylonurus ornatus Taxon name last Modified on 07/02/14	34	189	189	0	36	0	0	1
[107] † Laurieipterus elegans Taxon name last Modified on 07/02/14	94	129	129	0	36	0	0	1
[108] † Kokomopterus longicaudatus Taxon name last Modified on 07/02/14	44	178	178	0	37	0	0	1
[109] † Hardieopterus macrophthalmus Taxon name last Modified on 07/02/14	74	151	151	0	34	0	0	1
[110] † Drepanopterus pentlandicus Taxon name last Modified on 07/02/14	28	199	198	0	33	0	0	1
[111] † Hibbertopterus scouleri Taxon name last Modified on 07/02/14	31	196	196	0	32	0	0	1
[112] † Megarachne servinei Taxon name last Modified on 07/02/14	173	63	62	0	24	0	0	1
[113] † Chasmataspis laurencii Taxon name last Modified on 07/02/14	112	122	122	0	25	0	0	1
[114] † Loganamaraspis dunlopi Taxon name last Modified on 07/02/14	97	129	129	0	33	0	0	1
[115] † Achanarraspis reedi Taxon name last Modified on 07/02/14	153	86	86	0	20	0	0	1
[116] † Heteroaspis stoermeri Taxon name last Modified on 07/02/14	144	91	91	0	24	0	0	1
[117] † Diploaspis casteri Taxon name last Modified on 07/02/14	107	125	125	0	27	0	0	1
[118] † Diploaspis muelleri Taxon name last Modified on 07/02/14	148	87	87	0	24	0	0	1
[119] † Dvulikiaspis menneri Taxon name last Modified on 07/02/14	122	106	106	0	31	0	0	1
[120] † Octoberaspis ushakovi Taxon name last Modified on 07/02/14	65	160	160	0	34	0	0	1
[121] Centruroides vittatus Taxon name last Modified on 11/13/20	1	198	198	0	60	0	0	1
[122] Androctonus australis Taxon name last Modified on 07/02/14	0	198	198	0	61	0	0	1
[123] † Proscorpius osborni Taxon name last Modified on 07/02/14	29	169	169	0	61	0	0	1
[124] † Palaeophonus nuncius Taxon name last Modified on 07/02/14	54	146	146	0	59	0	0	1
[125] Caddo agilis Taxon name last Modified on 11/13/20	7	177	177	0	75	0	0	1
[126] Chileogovea oedipus Taxon name last Modified on 11/13/20	15	173	173	0	71	0	0	1
[127] Cyphophthalmus duricorius Taxon name last Modified on 07/02/14	15	173	173	0	71	0	0	1
[128] Gonyleptes horridus Taxon name last Modified on 07/02/14	7	178	178	0	74	0	0	1
[129] Leiobunum rotundum Taxon name last Modified on 07/02/14	7	177	177	0	75	0	0	1
[130] Sclerobunus robustus Taxon name last Modified on 07/02/14	7	178	178	0	74	0	0	1
[131] Allothrombium simoni Taxon name last Modified on 07/02/14	16	160	159	0	84	0	0	1
[132] Amblyomma americanum Taxon name last Modified on 11/13/20	14	158	157	0	88	0	0	1
[133] Neocarus texanus Taxon name last Modified on 07/02/14	8	167	167	0	84	0	0	1
[134] Cryptocellus goodnighti Taxon name last Modified on 07/02/14	4	180	180	0	75	0	0	1
[135] Poliochera punctulata Taxon name last Modified on 07/02/14	59	126	126	0	74	0	0	1
[136] Ricinoides atewa Taxon name last Modified on 07/02/14	4	180	180	0	75	0	0	1
[137] † Anthracomartus hindi Taxon name last Modified on 07/02/14	31	157	157	0	71	0	0	1
[138] † Eophrynus prestvicii Taxon name last Modified on 07/02/14	32	159	159	0	68	0	0	1
[139] † Gilboarachne griersoni Taxon name last Modified on 07/02/14	20	167	167	0	72	0	0	1
[140] † Palaeocharinus sp. Taxon name last Modified on 07/02/14	8	179	179	0	72	0	0	1
[141] † Mesotarbus peteri Taxon name last Modified on 07/02/14	51	136	136	0	72	0	0	1
[142] † Plesiosiro madeleyi Taxon name last Modified on 07/02/14	43	145	145	0	71	0	0	1
[143] Eukoenenia mirabilis Taxon name last Modified on 07/02/14	3	181	180	0	76	0	0	1
[144] Prokoenenia wheeleri Taxon name last Modified on 07/02/14	3	181	180	0	76	0	0	1
[145] Mastigoproctus giganteus Taxon name last Modified on 11/13/20	3	190	190	0	66	0	0	1
[146] Thelyphonus caudatus Taxon name last Modified on 07/02/14	3	186	186	0	70	0	0	1
[147] Hubbardia pentapeltis Taxon name last Modified on 07/02/14	3	186	185	0	71	0	0	1
[148] Schizomus crassicaudatus Taxon name last Modified on 07/02/14	3	186	185	0	71	0	0	1
[149] † Permarachne novokshonovi Taxon name last Modified on 07/02/14	119	101	101	0	39	0	0	1
[150] † Attercopus filmbriunguis Taxon name last Modified on 07/02/14	173	68	68	0	18	0	0	1
[151] Liphistius malayanus Taxon name last Modified on 07/02/14	3	175	175	0	81	0	0	1
[152] Aphonopelma hentzi Taxon name last Modified on 11/13/20	3	166	166	0	90	0	0	1
[153] Hypochilus pococki Taxon name last Modified on 11/13/20	3	166	166	0	90	0	0	1
[154] Cupienius getazi Taxon name last Modified on 07/02/14	4	165	165	0	90	0	0	1
[155] Neobisium maritimum Taxon name last Modified on 07/02/14	10	166	165	0	84	0	0	1
[156] Chelifer cancroides Taxon name last Modified on 11/13/20	10	165	164	0	85	0	0	1
[157] Eremocosta striata Taxon name last Modified on 07/02/14	10	162	161	0	88	0	0	1
[158] Galeodes armeniacus Taxon name last Modified on 07/02/14	12	161	160	0	87	0	0	1
[159] Phrynus marginemaculatus Taxon name last Modified on 07/02/14	4	182	182	0	73	0	0	1
[160] Charinus victori Taxon name last Modified on 07/02/14	4	181	181	0	74	0	0	1

Project downloads

type	number of downloads	Individual items downloaded (where applicable)
Total downloads from project	169
Project downloads	114
Matrix downloads	54	Xiph focus (54 downloads);
Media downloads	1	M736137 (1 download);