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the current taxonomic framework for Xiphosura – set out in the Treatise of Invertebrate Paleontology in 1955 – to be outdated and in need of revision, with several common genera such as Paleolimulus Dunbar, 1923 and Limulitella Størmer, 1952 acting as wastebasket taxon.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

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MorphoBank Project 3497

Creation Date:
11 July 2019
Publication Date:
13 November 2020
Project views: 14160
Media downloads: 1
Matrix downloads: 53

This research
supported by

Authors' Institutions

West Virginia University

Members

James Lamsdell
Project Administrator

160

259

27773
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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 Last Modified in 07/02/14	35	149	149	0	75	0	0	1
[2] † Leanchoilia illecebrosa Last Modified in 07/02/14	36	163	163	0	60	0	0	1
[3] † Alalcomenaeus cambricus Last Modified in 07/02/14	28	170	170	0	61	0	0	1
[4] † Yohoia tenuis Last Modified in 07/02/14	70	133	133	0	56	0	0	1
[5] † Emeraldella brocki Last Modified in 07/02/14	21	165	165	0	73	0	0	1
[6] † Sydneyia inexpectans Last Modified in 07/02/14	42	147	147	0	70	0	0	1
[7] † Olenoides serratus Last Modified in 07/02/14	13	188	188	0	58	0	0	1
[8] Pycnogonum litorale Last Modified in 07/02/14	8	102	102	0	149	0	0	1
[9] † Haliestes dasos Last Modified in 07/02/14	39	85	85	0	135	0	0	1
[10] † Palaeoisopus problematicus Last Modified in 07/02/14	22	116	116	0	121	0	0	1
[11] † Offacolus kingi Last Modified in 07/02/14	87	131	131	0	41	0	0	1
[12] † Dibasterium durgae Last Modified in 07/02/14	57	149	149	0	53	0	0	1
[13] † Anderella parva Last Modified in 08/20/14	156	87	87	0	16	0	0	1
[14] † Weinbergina opitzi Last Modified in 07/02/14	68	161	161	0	30	0	0	1
[15] † Camanchia grovensis Last Modified in 07/02/14	147	95	95	0	17	0	0	1
[16] † Venustulus waukeshaensis Last Modified in 07/02/14	128	111	111	0	20	0	0	1
[17] † Legrandella lombardi Last Modified in 07/02/14	137	107	107	0	15	0	0	1
[18] † Bembicosoma pomphicus Last Modified in 07/02/14	159	85	85	0	15	0	0	1
[19] † Bunodes lunula Last Modified in 11/13/20	156	86	86	0	17	0	0	1
[20] † Limuloides limuloides Last Modified in 07/02/14	148	97	97	0	14	0	0	1
[21] † Pasternakevia podolica Last Modified in 07/02/14	157	87	87	0	15	0	0	1
[22] † Cyamocephalus loganensis Last Modified in 07/02/14	150	94	94	0	15	0	0	1
[23] † Pseudoniscus roosevelti Last Modified in 07/02/14	156	88	88	0	15	0	0	1
[24] † Winneshiekia youngae Last Modified in 07/16/18	156	89	89	0	14	0	0	1
[25] † Houia yueya Last Modified in 05/25/15	144	89	89	0	26	0	0	1
[26] † Euproops anthrax Last Modified in 07/02/14	98	138	138	0	23	0	0	1
[27] † Euproops mariae Last Modified in 07/17/14	168	80	80	0	11	0	0	1
[28] † Euproops danae Last Modified in 07/02/14	93	143	143	0	23	0	0	1
[29] † Euproops rotundatus Last Modified in 07/17/14	146	101	101	0	12	0	0	1
[30] † Euproops sp Last Modified in 07/02/14	91	141	141	0	27	0	0	1
[31] † Bellinurus lunatus Last Modified in 07/18/14	148	98	98	0	13	0	0	1
[32] † Bellinurus arcuatus Last Modified in 07/18/14	149	97	97	0	13	0	0	1
[33] † Bellinurus reginae Last Modified in 07/18/14	149	96	96	0	14	0	0	1
[34] † Bellinurus truemanii Last Modified in 07/18/14	150	95	95	0	14	0	0	1
[35] † Bellinurus bellulus Last Modified in 07/18/14	153	92	92	0	14	0	0	1
[36] † Bellinurus trilobitoides Last Modified in 07/02/14	149	96	96	0	14	0	0	1
[37] † Anacontium carpenteri Last Modified in 07/16/14	220	37	37	0	2	0	0	1
[38] † Liomesaspis laevis Last Modified in 07/16/14	151	90	90	0	18	0	0	1
[39] † Stilpnocephalus pontebbanus Last Modified in 07/27/20	222	27	27	0	10	0	0	1
[40] † Alanops magnifica Last Modified in 07/16/14	59	157	157	0	43	0	0	1
[41] † Pringlia birtwelli Last Modified in 07/17/14	162	75	75	0	22	0	0	1
[42] † Kasibelinurus amoricum Last Modified in 07/02/14	154	93	93	0	12	0	0	1
[43] † Kasibelinurus randalli Last Modified in 07/02/14	164	83	83	0	12	0	0	1
[44] † Pickettia carteri Last Modified in 07/27/20	162	84	84	0	13	0	0	1
[45] † Lunataspis aurora Last Modified in 07/02/14	142	99	99	0	18	0	0	1
[46] Limulus polyphemus Last Modified in 07/02/14	0	219	219	0	40	0	0	1
[47] † Limulus darwini Last Modified in 07/27/20	162	80	80	0	17	0	0	1
[48] † Limulus coffini Last Modified in 07/14/14	199	46	46	0	14	0	0	1
[49] † Crenatolimulus paluxyensis Last Modified in 07/14/14	159	84	84	0	16	0	0	1
[50] Carcinoscorpius rotundicauda Last Modified in 07/03/14	0	220	220	0	39	0	0	1
[51] † Heterolimulus gadeai Last Modified in 07/16/18	149	94	94	0	16	0	0	1
[52] Tachypleus gigas Last Modified in 07/03/14	0	220	220	0	39	0	0	1
[53] † Tachypleus decheni Last Modified in 07/16/18	148	95	95	0	16	0	0	1
[54] † Tachypleus syriacus Last Modified in 07/16/18	143	101	100	0	16	0	0	1
[55] Tachypleus tridentatus Last Modified in 07/02/14	0	220	220	0	39	0	0	1
[56] † Yunnanolimulus luopingensis Last Modified in 07/11/14	30	189	189	0	40	0	0	1
[57] † Xaniopyramis linseyi Last Modified in 07/10/14	187	67	67	0	5	0	0	1
[58] † Moravurus rehori Last Modified in 07/27/20	231	23	23	0	5	0	0	1
[59] † Paleolimulus woodae Last Modified in 07/11/18	157	89	89	0	13	0	0	1
[60] † Paleolimulus signatus Last Modified in 07/02/14	46	183	183	0	30	0	0	1
[61] † Paleolimulus kunguricus Last Modified in 07/16/18	141	104	104	0	14	0	0	1
[62] † Bellinuroopsis rossicus Last Modified in 07/18/14	152	93	93	0	14	0	0	1
[63] † Rolfeia fouldenensis Last Modified in 07/10/14	143	102	102	0	14	0	0	1
[64] † Tarracolimulus rieki Last Modified in 07/14/14	155	89	89	0	15	0	0	1
[65] † Casterolimulus kletti Last Modified in 07/16/14	221	36	36	0	2	0	0	1
[66] † Limulitella vicensis Last Modified in 07/21/14	175	72	72	0	12	0	0	1
[67] † Victalimulus mcqueeni Last Modified in 07/16/14	116	125	125	0	18	0	0	1
[68] † Limulus woodwardi Last Modified in 07/13/18	222	35	35	0	2	0	0	1
[69] † Mesolimulus crespelli Last Modified in 07/15/14	164	79	79	0	16	0	0	1
[70] † Mesolimulus tafraoutensis Last Modified in 07/27/20	155	88	88	0	16	0	0	1
[71] † Mesolimulus walchi Last Modified in 07/02/14	57	173	173	0	29	0	0	1
[72] † Mesolimulus sibiricus Last Modified in 07/27/20	161	83	83	0	15	0	0	1
[73] † Panduralimulus babcocki Last Modified in 07/11/14	142	98	98	0	19	0	0	1
[74] † Tasmaniolimulus patersoni Last Modified in 07/27/20	160	80	80	0	19	0	0	1
[75] † Austrolimulus fletcheri Last Modified in 07/11/14	149	91	91	0	19	0	0	1
[76] † Vaderlimulus tricki Last Modified in 07/11/18	147	92	92	0	20	0	0	1
[77] † Psammolimulus gottingensis Last Modified in 07/11/14	113	122	122	0	24	0	0	1
[78] † Limulitella bronni Last Modified in 07/17/14	149	90	90	0	20	0	0	1
[79] † Limulitella tejraensis Last Modified in 07/11/18	160	79	79	0	20	0	0	1
[80] † Limulitella henkeli Last Modified in 07/17/14	169	75	75	0	15	0	0	1
[81] † Madagascan limulid Last Modified in 07/23/19	165	79	79	0	15	0	0	1
[82] † Paleolimulus fuchsbergensis Last Modified in 07/11/14	156	84	84	0	19	0	0	1
[83] † Dubbolimulus peetae Last Modified in 07/16/18	146	93	93	0	20	0	0	1
[84] † Paleolimulus longispinus Last Modified in 07/15/14	137	100	100	0	22	0	0	1
[85] † Paleolimulus jakovlevi Last Modified in 07/27/20	165	78	78	0	16	0	0	1
[86] † Valloisella lievinensis Last Modified in 07/16/14	146	97	97	0	16	0	0	1
[87] † Stoermeropterus conicus Last Modified in 07/02/14	25	199	199	0	35	0	0	1
[88] † Moselopterus ancylotelson Last Modified in 07/02/14	24	195	195	0	40	0	0	1
[89] † Onychopterella kokomoensis Last Modified in 07/02/14	33	190	190	0	36	0	0	1
[90] † Eurypterus tetragonophthalmus Last Modified in 07/02/14	10	214	214	0	35	0	0	1
[91] † Erieopterus macrophthalmus Last Modified in 07/02/14	33	191	191	0	35	0	0	1
[92] † Strobilopterus proteus Last Modified in 07/02/14	47	188	188	0	24	0	0	1
[93] † Dolichopterus macrocheirus Last Modified in 07/02/14	23	202	202	0	34	0	0	1
[94] † Carcinosoma newlini Last Modified in 07/02/14	31	189	189	0	39	0	0	1
[95] † Megalograptus ohioensis Last Modified in 07/02/14	22	202	202	0	35	0	0	1
[96] † Mixopterus kiaeri Last Modified in 07/02/14	32	188	188	0	39	0	0	1
[97] † Orcanopterus manitoulinensis Last Modified in 07/02/14	73	146	146	0	40	0	0	1
[98] † Adelophthalmus sievertsi Last Modified in 07/02/14	46	176	176	0	37	0	0	1
[99] † Nanahughmilleria norvegica Last Modified in 07/02/14	24	199	199	0	36	0	0	1
[100] † Hughmilleria socialis Last Modified in 07/02/14	23	196	196	0	40	0	0	1
[101] † Slimonia acuminata Last Modified in 07/02/14	25	194	194	0	40	0	0	1
[102] † Erettopterus bilobus Last Modified in 07/02/14	25	192	192	0	42	0	0	1
[103] † Pterygotus anglicus Last Modified in 07/02/14	31	185	185	0	43	0	0	1
[104] † Brachyopterus stubblefieldi Last Modified in 07/02/14	71	151	151	0	37	0	0	1
[105] † Rhenopterus diensti Last Modified in 07/02/14	24	199	199	0	36	0	0	1
[106] † Parastylonurus ornatus Last Modified in 07/02/14	34	189	189	0	36	0	0	1
[107] † Laurieipterus elegans Last Modified in 07/02/14	94	129	129	0	36	0	0	1
[108] † Kokomopterus longicaudatus Last Modified in 07/02/14	44	178	178	0	37	0	0	1
[109] † Hardieopterus macrophthalmus Last Modified in 07/02/14	74	151	151	0	34	0	0	1
[110] † Drepanopterus pentlandicus Last Modified in 07/02/14	28	199	198	0	33	0	0	1
[111] † Hibbertopterus scouleri Last Modified in 07/02/14	31	196	196	0	32	0	0	1
[112] † Megarachne servinei Last Modified in 07/02/14	173	63	62	0	24	0	0	1
[113] † Chasmataspis laurencii Last Modified in 07/02/14	112	122	122	0	25	0	0	1
[114] † Loganamaraspis dunlopi Last Modified in 07/02/14	97	129	129	0	33	0	0	1
[115] † Achanarraspis reedi Last Modified in 07/02/14	153	86	86	0	20	0	0	1
[116] † Heteroaspis stoermeri Last Modified in 07/02/14	144	91	91	0	24	0	0	1
[117] † Diploaspis casteri Last Modified in 07/02/14	107	125	125	0	27	0	0	1
[118] † Diploaspis muelleri Last Modified in 07/02/14	148	87	87	0	24	0	0	1
[119] † Dvulikiaspis menneri Last Modified in 07/02/14	122	106	106	0	31	0	0	1
[120] † Octoberaspis ushakovi Last Modified in 07/02/14	65	160	160	0	34	0	0	1
[121] Centruroides vittatus Last Modified in 11/13/20	1	198	198	0	60	0	0	1
[122] Androctonus australis Last Modified in 07/02/14	0	198	198	0	61	0	0	1
[123] † Proscorpius osborni Last Modified in 07/02/14	29	169	169	0	61	0	0	1
[124] † Palaeophonus nuncius Last Modified in 07/02/14	54	146	146	0	59	0	0	1
[125] Caddo agilis Last Modified in 11/13/20	7	177	177	0	75	0	0	1
[126] Chileogovea oedipus Last Modified in 11/13/20	15	173	173	0	71	0	0	1
[127] Cyphophthalmus duricorius Last Modified in 07/02/14	15	173	173	0	71	0	0	1
[128] Gonyleptes horridus Last Modified in 07/02/14	7	178	178	0	74	0	0	1
[129] Leiobunum rotundum Last Modified in 07/02/14	7	177	177	0	75	0	0	1
[130] Sclerobunus robustus Last Modified in 07/02/14	7	178	178	0	74	0	0	1
[131] Allothrombium simoni Last Modified in 07/02/14	16	160	159	0	84	0	0	1
[132] Amblyomma americanum Last Modified in 11/13/20	14	158	157	0	88	0	0	1
[133] Neocarus texanus Last Modified in 07/02/14	8	167	167	0	84	0	0	1
[134] Cryptocellus goodnighti Last Modified in 07/02/14	4	180	180	0	75	0	0	1
[135] Poliochera punctulata Last Modified in 07/02/14	59	126	126	0	74	0	0	1
[136] Ricinoides atewa Last Modified in 07/02/14	4	180	180	0	75	0	0	1
[137] † Anthracomartus hindi Last Modified in 07/02/14	31	157	157	0	71	0	0	1
[138] † Eophrynus prestvicii Last Modified in 07/02/14	32	159	159	0	68	0	0	1
[139] † Gilboarachne griersoni Last Modified in 07/02/14	20	167	167	0	72	0	0	1
[140] † Palaeocharinus sp. Last Modified in 07/02/14	8	179	179	0	72	0	0	1
[141] † Mesotarbus peteri Last Modified in 07/02/14	51	136	136	0	72	0	0	1
[142] † Plesiosiro madeleyi Last Modified in 07/02/14	43	145	145	0	71	0	0	1
[143] Eukoenenia mirabilis Last Modified in 07/02/14	3	181	180	0	76	0	0	1
[144] Prokoenenia wheeleri Last Modified in 07/02/14	3	181	180	0	76	0	0	1
[145] Mastigoproctus giganteus Last Modified in 11/13/20	3	190	190	0	66	0	0	1
[146] Thelyphonus caudatus Last Modified in 07/02/14	3	186	186	0	70	0	0	1
[147] Hubbardia pentapeltis Last Modified in 07/02/14	3	186	185	0	71	0	0	1
[148] Schizomus crassicaudatus Last Modified in 07/02/14	3	186	185	0	71	0	0	1
[149] † Permarachne novokshonovi Last Modified in 07/02/14	119	101	101	0	39	0	0	1
[150] † Attercopus filmbriunguis Last Modified in 07/02/14	173	68	68	0	18	0	0	1
[151] Liphistius malayanus Last Modified in 07/02/14	3	175	175	0	81	0	0	1
[152] Aphonopelma hentzi Last Modified in 11/13/20	3	166	166	0	90	0	0	1
[153] Hypochilus pococki Last Modified in 11/13/20	3	166	166	0	90	0	0	1
[154] Cupienius getazi Last Modified in 07/02/14	4	165	165	0	90	0	0	1
[155] Neobisium maritimum Last Modified in 07/02/14	10	166	165	0	84	0	0	1
[156] Chelifer cancroides Last Modified in 11/13/20	10	165	164	0	85	0	0	1
[157] Eremocosta striata Last Modified in 07/02/14	10	162	161	0	88	0	0	1
[158] Galeodes armeniacus Last Modified in 07/02/14	12	161	160	0	87	0	0	1
[159] Phrynus marginemaculatus Last Modified in 07/02/14	4	182	182	0	73	0	0	1
[160] Charinus victori Last Modified in 07/02/14	4	181	181	0	74	0	0	1

Project views

type	number of views	Individual items viewed (where applicable)
Total project views	14160
Project overview	1366
Matrix views	796	Matrix landing page (666 views); Xiph focus (130 views);
Media views	2099	Media search (1801 views); M736137 (298 views);
Views for media list	662
Bibliography	454
Specimen list	1785
Taxon list	6992
Documents list	6

Project downloads

type	number of downloads	Individual items downloaded (where applicable)
Total downloads from project	167
Project downloads	113
Matrix downloads	53	Xiph focus (53 downloads);
Media downloads	1	M736137 (1 download);