Morphobank

Project 2084: J. D. Carrillo, R. J. Asher. 2017. An exceptionally well preserved skeleton Thomashuxleya externa (Mammalia, Notoungulata), from the Eocene of Patagonia, Argentina. Palaeontologia electronica. 20.2.34A:1-33.

Specimen: † Thomashuxleya externa (MEPF-PV 8166)
View: Artistic reconstruction

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Abstract

We describe one of the oldest notoungulate skeletons with associated craniodental and postcranial elements: Thomashuxleya externa (Isotemnidae) from Cañadón Vaca in Patagonia, Argentina (Vacan subage of the Casamayoran SALMA, middle Eocene). We provide body mass estimates given by different elements of the skeleton, describe the bone histology, and study its phylogenetic position. We note differences in the scapulae, humerii, ulnae and radii of the new specimen in comparison with other specimens previously referred to this taxon. We estimate a body mass of 84 ± 24.2 kg, showing that notoungulates had acquired a large body mass by the middle Eocene. Bone histology shows that the new specimen was skeletally mature. The new material supports the placement of Thomashuxleya as an early, divergent member of Toxodontia. Among placentals, our phylogenetic analysis of a combined DNA, collagen, and morphology matrix favor only a limited number of possible phylogenetic relationships, but cannot yet arbitrate between potential affinities with Afrotheria or Laurasiatheria. With no constraint, maximum parsimony supports Thomashuxleya and Carodnia with Afrotheria. With Notoungulata and Litopterna constrained as monophyletic (including Macrauchenia and Toxodon known for collagens), these clades are reconstructed on the stem to Euungulata (i.e., Perissodactyla and Artiodactyla). Unconstrained, Bayesian analysis weakly supports the possibility that Thomashuxleya is a stem xenarthran; with Notoungulata and Litopterna constrained as monophyletic, the two clades are recovered as sister to Perissodactyla. Anatomical data sampled thus far for Thomashuxleya, combined with collagen amino acids for Pleistocene meridiungulates, substantially limit the number of possible affinities for endemic South American species among mammals, although ambiguity still remains.

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Project DOI: 10.7934/P2084, http://dx.doi.org/10.7934/P2084

This project contains	Matrices
1 Media 2 Matrices 6 Documents 154 Taxa 1 Specimen 3806 Characters Total size of project's media files: 8.04M Download Project SDD File	Total scored cells: 257229 Total media associated with cells: 0 Total labels associated with cell media: 0
	Characters
	Total characters: 3806 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 8991 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:3792/14

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MorphoBank Project 2084

Creation Date:
15 December 2014
Publication Date:
19 July 2017
Project views: 25567
Media downloads: 2
Matrix downloads: 79

Authors' Institutions

Paläontologisches Museum, Zürich

University of Cambridge

University of Zurich (Universität Zürich)

Members

Juan David Carrillo
Project Administrator

154

3806

257229
(175656, 0, 81573)

Robert Asher
Full membership

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Taxonomic Overview for Matrix 'M22531' (86 Taxa)

taxon	unscored cells	scored cells	no cell support	NPA cells	"-" cells	cell images	labels on cell images	member access
[1] † Thomashuxleya externa Last Modified in 07/19/17	1653	1500	1486	0	521	0	0	2
[2] † Carodnia vieirai Last Modified in 07/19/17	1702	1464	464	0	494	0	0	2
[3] † Protolipterna ellipsodontoides Last Modified in 07/19/17	2366	981	332	0	314	0	0	2
[4] † Didolodus multicuspis Last Modified in 07/19/17	2482	891	891	0	287	0	0	2
[5] † Morganucodon oehleri Last Modified in 07/19/17	1303	1328	1285	0	1032	0	0	2
[6] † Morganucodon watsoni Last Modified in 07/19/17	747	1720	1716	0	1196	0	0	2
[7] † Haldanodon exspectatus Last Modified in 07/19/17	1073	1601	1573	0	992	0	0	2
[8] Ornithorhynchus anatinus Last Modified in 12/16/14	53	1930	1895	0	1682	0	0	2
[9] Tachyglossus aculeatus Last Modified in 12/16/14	19	1546	1344	0	2099	0	0	2
[10] † Zhangheotherium quinquecuspidens Last Modified in 07/19/17	816	1609	1586	0	1238	0	0	2
[11] † Henkelotherium guimarotae Last Modified in 07/19/17	1872	1083	1083	0	705	0	0	2
[12] † Pucadelphys andinus Last Modified in 07/19/17	800	2066	2030	0	801	0	0	2
[13] Didelphis virginiana Last Modified in 12/16/14	11	2743	593	0	922	0	0	2
[14] Dromiciops gliroides Last Modified in 12/16/14	302	2482	562	0	882	0	0	2
[15] † Eomaia scansoria Last Modified in 07/19/17	1835	1413	1410	0	412	0	0	2
[16] † Maelestes gobiensis Last Modified in 07/19/17	1690	1454	1229	0	518	0	0	2
[17] Dasypus novemcinctus Last Modified in 12/16/14	34	1836	600	0	1804	0	0	2
[18] † Hapalops elongatus Last Modified in 07/19/17	463	1519	10	0	1706	0	0	2
[19] Tamandua tetradactyla Last Modified in 12/16/14	24	1729	83	0	1917	0	0	2
[20] Manis pentadactyla Last Modified in 12/16/14	60	1590	785	0	2019	0	0	2
[21] † Metacheiromys marshi Last Modified in 07/19/17	1912	1424	2	0	324	0	0	2
[22] † Ukhaatherium nessovi Last Modified in 07/19/17	1650	1481	1358	0	530	0	0	2
[23] † Zalambdalestes lechei Last Modified in 07/19/17	1463	1677	1637	0	526	0	0	2
[24] † Leptictis dakotensis Last Modified in 07/19/17	534	2494	2457	0	634	0	0	2
[25] Solenodon paradoxus Last Modified in 12/16/14	31	2758	2373	0	891	0	0	2
[26] Talpa europaea Last Modified in 12/16/14	89	2583	2445	0	1000	0	0	2
[27] Erinaceus europaeus Last Modified in 12/16/14	37	2665	2630	0	968	0	0	2
[28] Sorex araneus Last Modified in 12/16/14	525	1941	1896	0	1197	0	0	2
[29] Amblysomus hottentotus Last Modified in 12/16/14	354	2332	2004	0	987	0	0	2
[30] Echinops telfairi Last Modified in 12/16/14	314	2359	2209	0	993	0	0	2
[31] Rhynchocyon cirnei Last Modified in 12/16/14	50	2708	2542	0	910	0	0	2
[32] Procavia capensis Last Modified in 07/19/17	51	2807	2487	0	828	0	0	2
[33] Loxodonta africana Last Modified in 12/16/14	69	2194	2124	0	1398	0	0	2
[34] Trichechus manatus Last Modified in 12/16/14	193	1820	1765	0	1660	0	0	2
[35] † Moeritherium lyonsi Last Modified in 07/19/17	1453	1543	1452	0	665	0	0	2
[36] Galeopterus variegatus Last Modified in 12/16/14	348	2555	2319	0	762	0	0	2
[37] Cynocephalus volans Last Modified in 12/16/14	94	2780	15	0	796	0	0	2
[38] Ptilocercus lowii Last Modified in 12/16/14	330	2693	770	0	642	0	0	2
[39] Tupaia glis Last Modified in 12/16/14	41	2913	925	0	713	0	0	2
[40] Homo sapiens Last Modified in 12/16/14	29	2762	2509	0	932	0	0	2
[41] Lemur catta catta Last Modified in 12/16/14	115	2808	721	0	742	0	0	2
[42] Saimiri sciureus Last Modified in 12/16/14	88	2717	768	0	863	0	0	2
[43] Tarsius syrichta Last Modified in 12/16/14	277	2472	172	0	912	0	0	2
[44] † Notharctus tenebrosus Last Modified in 07/19/17	626	2420	1142	0	620	0	0	2
[45] † Protungulatum donnae Last Modified in 07/19/17	2345	1009	987	0	306	0	0	2
[46] † Apheliscus insidiosus Last Modified in 07/19/17	2229	1027	417	0	404	0	0	2
[47] † Phenacodus intermedius Last Modified in 07/19/17	502	2535	1420	0	624	0	0	2
[48] † Hyopsodus paulus Last Modified in 07/19/17	1018	1996	1992	0	650	0	0	2
[49] Felis silvestris Last Modified in 12/16/14	17	2521	2511	0	1127	0	0	2
[50] Canis lupus familiaris Last Modified in 12/16/14	19	2879	2863	0	767	0	0	2
[51] † Vulpavus ovatus Last Modified in 07/19/17	1545	1625	2	0	493	0	0	2
[52] † Vulpavus profectus Last Modified in 07/19/17	1233	1890	2	0	540	0	0	2
[53] † Sinopa rapax Last Modified in 07/19/17	1318	1850	1842	0	496	0	0	2
[54] Oryctolagus cuniculus Last Modified in 12/16/14	128	2431	2425	0	1102	0	0	2
[55] † Dawsonolagus antiquus Last Modified in 07/19/17	2514	610	607	0	536	0	0	2
[56] † Gomphos elkema Last Modified in 07/19/17	910	1892	1	0	858	0	0	2
[57] † Rhombomylus turpanensis Last Modified in 07/19/17	550	2132	2103	0	996	0	0	2
[58] † Tribosphenomys minutus Last Modified in 07/19/17	1991	996	977	0	685	0	0	2
[59] † Paramys delicatus Last Modified in 07/19/17	676	1975	1665	0	1021	0	0	2
[60] † Cocomys lingchaensis Last Modified in 07/19/17	1376	1376	1353	0	917	0	0	2
[61] Ictidomys tridecemlineatus Last Modified in 12/16/14	300	2293	2211	0	1073	0	0	2
[62] Castor canadensis Last Modified in 12/16/14	79	2414	2319	0	1188	0	0	2
[63] Cavia porcellus Last Modified in 12/16/14	13	2246	2196	0	1420	0	0	2
[64] Rattus norvegicus Last Modified in 12/16/14	4	2201	1886	0	1466	0	0	2
[65] Orycteropus afer Last Modified in 12/16/14	296	1867	1400	0	1502	0	0	2
[66] Equus caballus Last Modified in 12/16/14	78	2709	2178	0	883	0	0	2
[67] Sus scrofa Last Modified in 12/16/14	204	2617	2017	0	854	0	0	2
[68] Tursiops truncatus Last Modified in 12/16/14	95	1163	836	0	2410	0	0	2
[69] Hippopotamus amphibius Last Modified in 12/16/14	174	2549	2181	0	942	0	0	2
[70] Lama glama Last Modified in 12/16/14	161	2428	2034	0	1074	0	0	2
[71] Bos taurus Last Modified in 12/16/14	77	2579	2173	0	1010	0	0	2
[72] † Mesohippus bairdi Last Modified in 07/19/17	740	2242	1939	0	682	0	0	2
[73] Caperea marginata Last Modified in 12/16/14	348	834	701	0	2486	0	0	2
[74] † Basilosaurus cetoides Last Modified in 07/19/17	1703	1040	815	0	917	0	0	2
[75] † Mesonyx obtusidens Last Modified in 07/19/17	1093	1729	1447	0	838	0	0	2
[76] † Archaeotherium mortoni Last Modified in 07/19/17	839	1959	1754	0	864	0	0	2
[77] † Artiocetus clavis Last Modified in 07/19/17	2450	750	679	0	461	0	0	2
[78] † Rodhocetus balochistanensis Last Modified in 07/19/17	3196	386	339	0	78	0	0	2
[79] Pteropus giganteus Last Modified in 12/16/14	51	2165	1072	0	1445	0	0	2
[80] Rhinopoma hardwickii Last Modified in 12/16/14	347	2342	1291	0	974	0	0	2
[81] Saccopteryx bilineata Last Modified in 12/16/14	356	2353	1399	0	952	0	0	2
[82] Nycteris thebaica Last Modified in 12/16/14	495	2163	1418	0	1002	0	0	2
[83] Myotis lucifugus Last Modified in 12/16/14	340	2257	1523	0	1065	0	0	2
[84] Pteronotus parnellii Last Modified in 12/16/14	445	2351	1687	0	868	0	0	2
[85] † Icaronycteris index Last Modified in 07/19/17	1297	1818	1135	0	549	0	0	2
[86] † Onychonycteris finneyi Last Modified in 07/19/17	1715	1471	1002	0	486	0	0	2

Taxonomic Overview for Matrix 'M24667' (68 Taxa)

taxon	unscored cells	scored cells	no cell support	NPA cells	"-" cells	cell images	labels on cell images	member access
[1] † Leptictis Last Modified in 07/19/17	8	108	108	0	30	0	0	2
[2] † Phenacodus Last Modified in 07/19/17	7	102	102	0	37	0	0	2
[3] † Meniscotherium Last Modified in 07/19/17	5	105	105	0	36	0	0	2
[4] † Protolipterna Last Modified in 07/19/17	102	23	23	0	21	0	0	2
[5] † Proterotherium Last Modified in 07/19/17	9	93	93	0	44	0	0	2
[6] † Trigonostylops Last Modified in 07/19/17	19	95	95	0	32	0	0	2
[7] † Astrapotherium Last Modified in 07/19/17	9	99	98	0	39	0	0	2
[8] † Pyrotherium Last Modified in 07/19/17	42	79	79	0	25	0	0	2
[9] † Notostylops Last Modified in 07/19/17	4	138	138	0	4	0	0	2
[10] † Henricosbornia Last Modified in 07/19/17	101	43	43	0	2	0	0	2
[11] † Simpsonotus Last Modified in 07/19/17	25	119	119	0	2	0	0	2
[12] † Pleurostylodon Last Modified in 07/19/17	15	130	129	0	2	0	0	2
[13] † Thomashuxleya Last Modified in 07/19/17	35	109	109	0	2	0	0	2
[14] † Pampahippus Last Modified in 07/19/17	81	64	59	0	2	0	0	2
[15] † Periphragnis Last Modified in 07/19/17	9	136	135	0	2	0	0	2
[16] † Ryphodon Last Modified in 07/19/17	54	89	88	0	3	0	0	2
[17] † Puelia Last Modified in 07/19/17	54	91	90	0	1	0	0	2
[18] † Leontinia Last Modified in 07/19/17	16	127	127	0	3	0	0	2
[19] † Colpodon Last Modified in 07/19/17	24	119	119	0	3	0	0	2
[20] † Ancylocoelus Last Modified in 07/19/17	42	101	100	0	4	0	0	2
[21] † Scarittia Last Modified in 07/19/17	34	110	110	0	2	0	0	2
[22] † Homalodotherium Last Modified in 07/19/17	15	128	128	0	3	0	0	2
[23] † Asmodeus Last Modified in 07/19/17	68	75	75	0	3	0	0	2
[24] † Rhynchippus Last Modified in 07/19/17	16	129	129	0	1	0	0	2
[25] † Eurygenium Last Modified in 07/19/17	60	85	84	0	1	0	0	2
[26] † Pascualihippus Last Modified in 07/19/17	63	83	82	0	1	0	0	2
[27] † Eomorphippus Last Modified in 07/19/17	73	74	72	0	1	0	0	2
[28] † Morphippus Last Modified in 07/19/17	35	110	110	0	1	0	0	2
[29] † Argyrohippus Last Modified in 07/19/17	33	113	112	0	1	0	0	2
[30] † Nesodon Last Modified in 07/19/17	13	131	129	0	3	0	0	2
[31] † Adinotherium Last Modified in 07/19/17	11	133	132	0	3	0	0	2
[32] † Hoffstetterius Last Modified in 07/19/17	41	98	98	0	7	0	0	2
[33] † Ponanskytherium Last Modified in 07/19/17	38	100	100	0	8	0	0	2
[34] † Toxodon Last Modified in 07/19/17	40	98	98	0	8	0	0	2
[35] † Colbertia Last Modified in 07/19/17	29	117	116	0	1	0	0	2
[36] † Oldfieldthomasia Last Modified in 07/19/17	16	128	128	0	2	0	0	2
[37] † Campanorco Last Modified in 07/19/17	50	94	94	0	2	0	0	2
[38] † Acropithecus Last Modified in 07/19/17	73	71	71	0	2	0	0	2
[39] † Ultrapithecus Last Modified in 07/19/17	101	43	43	0	2	0	0	2
[40] † Notopithecus Last Modified in 07/19/17	9	135	133	0	4	0	0	2
[41] † Archaeophylus Last Modified in 07/19/17	125	18	18	0	3	0	0	2
[42] † Cochilius Last Modified in 07/19/17	46	97	97	0	3	0	0	2
[43] † Plagiarthrus Last Modified in 07/19/17	54	87	87	0	5	0	0	2
[44] † Protypotherium Last Modified in 07/19/17	22	120	119	0	5	0	0	2
[45] † Miocochilius Last Modified in 07/19/17	52	89	89	0	5	0	0	2
[46] † Interatherium Last Modified in 07/19/17	36	104	104	0	6	0	0	2
[47] † Federicoanaya Last Modified in 07/19/17	41	100	100	0	5	0	0	2
[48] † Trachytherus Last Modified in 07/19/17	16	128	128	0	2	0	0	2
[49] † Plesiotypotherium Last Modified in 07/19/17	29	104	104	0	13	0	0	2
[50] † Mesotherium Last Modified in 07/19/17	33	100	100	0	13	0	0	2
[51] † Eohyrax Last Modified in 07/19/17	92	52	52	0	2	0	0	2
[52] † Pseudhyrax Last Modified in 07/19/17	77	67	67	0	2	0	0	2
[53] † Archaeotypotherium Last Modified in 07/19/17	64	80	80	0	2	0	0	2
[54] † Archaeohyrax Last Modified in 07/19/17	14	131	130	0	2	0	0	2
[55] † Prohegetotherium Last Modified in 07/19/17	58	86	86	0	2	0	0	2
[56] † Hegetotherium Last Modified in 07/19/17	32	113	112	0	2	0	0	2
[57] † Prosotherium Last Modified in 07/19/17	56	80	80	0	10	0	0	2
[58] † Paedotherium Last Modified in 07/19/17	43	90	90	0	13	0	0	2
[59] † Zalambdalestes Last Modified in 07/19/17	2	103	103	0	41	0	0	2
[60] † Macrauchenia Last Modified in 07/19/17	13	101	101	0	32	0	0	2
[61] † Pampahippus secundus Last Modified in 07/19/17	71	76	74	0	1	0	0	2
[62] † Plexotemnus Last Modified in 07/19/17	101	45	44	0	1	0	0	2
[63] † Coquenia Last Modified in 07/19/17	10	135	135	0	1	0	0	2
[64] † Griphotherion Last Modified in 07/19/17	21	123	123	0	2	0	0	2
[65] † Boreastylops Last Modified in 07/19/17	22	120	120	0	4	0	0	2
[66] † Pampatemnus deuteros Last Modified in 07/19/17	39	107	107	0	0	0	0	2
[67] † Pampatemnus infernalis Last Modified in 07/19/17	76	69	69	0	1	0	0	2
[68] † Martinmiguelia fernandezi Last Modified in 07/19/17	101	45	45	0	0	0	0	2

Project views

type	number of views	Individual items viewed (where applicable)
Total project views	25567
Media views	3373	Media search (2888 views); M425185 (485 views);
Project overview	3255
Documents list	931
Views for media list	1063
Specimen list	1693
Taxon list	12618
Bibliography	820
Matrix views	1814	Matrix landing page (1283 views); Relationships of Thomashuxleya within Notoungulata (149 views); Relationships of Thomashuxleya within Placentals (382 views);

Project downloads

type	number of downloads	Individual items downloaded (where applicable)
Total downloads from project	431
Document downloads	8	Matrix with constraint (1 download); Matrix with no constraints for Notoungulata (3 downloads); Matrix with constraint for Notoungulata and Litopterna (3 downloads); Notoungulata data matrix (1 download);
Matrix downloads	79	Relationships of Thomashuxleya within Placentals (60 downloads); Relationships of Thomashuxleya within Notoungulata (19 downloads);
Project downloads	342
Media downloads	2	M425185 (2 downloads);