identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E487FEFFCAFF85CCDC9A30ACE3276B.text	03E487FEFFCAFF85CCDC9A30ACE3276B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Macraucheniidae	<div><p>Macraucheniidae (Figs 4A, D–E, 5A–D)</p><p>With 18 currently accepted genera,  Macraucheniidae is the second most diverse family of litopterns (Supporting information, Table S1), being particularly diverse during the Neogene alongside proterotheriids (Fig. 2B–C). It was first proposed by Gervais (1855) as a member of the order  Perissodactyla (‘Ordre des Jumentés’) to include one species,  Macrauchenia patachonica Owen, 1838 . This taxon was discovered by Charles Darwin in 1834 (Keynes 2001) and later described by Owen (1838), being the first litoptern known to the scientific community. Later, Ameghino (1902a), based mostly on the position of the external nares in the cranium, subdivided this family into three subfamilies:  Cramaucheniinae represented by  Cramauchenia Ameghino 1902a from Deseadean to Colhuehuapian South American Land Mammal Ages [hereafter SALMAs; Stages of Cione and Tonni (1995)], Theosodontinae represented by  Theosodon Ameghino, 1887 (Colhuehuapian to Laventan SALMAs), and  Macraucheniinae represented by  Macrauchenia (Pliocene to Holocene). Simpson (1945) reclassified  Macraucheniidae into the subfamilies  Macraucheniinae and  Adianthinae, without giving an anatomical justification for this change (Table 1). By Simpson’s (1945) definition,  Macraucheniinae was composed of the members of the three subfamilies mentioned by Ameghino (1902a) and close Neogene relatives discovered since then (e.g.,  Promacrauchenia Ameghino, 1904a), but also included  Paramacrauchenia Bordas, 1939 and  Victorlemoinea Ameghino, 1901, the former now considered a proterotheriid and the latter a sparnotheriodontid (Soria 2001; see  Sparnotheriodontidae section for more details). The subfamily  Adianthinae was composed of adianthids such as  Adianthus (see  Adianthidae section for more details).</p><p>Similarly to Ameghino (1902a), Soria (1981) recognized important anatomical differences between macraucheniids before and after the Chasicoan SALMA (Late Miocene) that justified a subfamilial division, separating  Macraucheniidae into two subfamilies:  Cramaucheniinae (pre-Chasicoan SALMA) and  Macraucheniinae (Chasicoan and post-Chasicoan SALMA or post-Santacrucian SALMA). This separation was based on the fact that  Macraucheniinae tends to have more derived features than  Cramaucheniinae, possessing a retracted nasal aperture to a more centrodorsal position and fused zeugopodial elements in the forelimbs (ulna-radius) and hind limbs (tibia-fibula; Soria 1981). However, of these two subfamilies only  Macraucheniinae is monophyletic according to phylogenetic analyses (Schmidt and Ferrero 2014, Forasiepi et al. 2016, McGrath et al. 2018, Püschel et al. 2023; Table 2).</p><p>Macraucheniids are distinguished from the two other families,  Adianthidae and  Proterotheriidae . Among other differences, macraucheniids lack the trilobed m3, which is characteristic of adianthids (Cifelli and Soria 1983a). Also, macraucheniids are functional tridactyls,unlike proterotheriids that show reduced(or lost) lateral digits II and IV (Soria 2001). However, the origin of  Macraucheniidae in the Palaeogene and the affinities of the family within  Litopterna are still contentious topics. The oldest uncontroversial members of the family are  Coniopternium Ameghino 1894a,  Cramauchenia Ameghino 1902a, and  Pternoconius Cifelli and Soria 1983b, all with a first appearance datum (FAD) in the Deseadean SALMA (Late Oligocene; Dozo and Vera 2010). These genera show dental and/or postcranial anatomical features that link them closely to later macraucheniids, like the presence of a mesolophid or cristid that connects the cristid obliqua with the entoconid [=entolophid (Soria and Hoffstetter 1985)] in all or some of the lower molars. Before the Oligocene,  Polymorphis Roth, 1899 from the Late Eocene of Patagonia has been proposed as the earliest member of  Macraucheniidae (Soria 1982, Cifelli 1983a), revalidating a previous proposal of Ameghino (1904b). Cifelli (1983a) even considered it as a member of a different subfamily,  Polymorphinae, within  Macraucheniidae (Table 1). Previously,  Polymorphinae was also considered a distinct family of the  Proterotheriidae (Odreman Rivas 1969) . Key features for associating  Polymorphis with macraucheniids are related to similarities in the upper molars, in particular, the presence of an oblique crest that connects the protocone with the metacone (Soria 1982; Fig. 4A; see also  Polymorphis spp. in Supporting information, File S2). In contrast, the lower molars of  Polymorphis spp. show important differences with Deseadean SALMA macraucheniids, such as the complete absence of a mesolophid. The proposal of  Polymorphis as a macraucheniid found support in a phylogenetic analysis (Cifelli 1993; Fig. 1C), and since then most recent studies have considered this taxon as the earliest member of  Macraucheniidae without any further testing (e.g., Dozo and Vera 2010, Croft et al. 2020; Table 2). Cifelli (1993) also found  Macraucheniidae as the sister group of  Adianthidae (Macrauchenioidea), and both families closely related to  Proterotheriidae ( Lopholipterna; Fig. 1C; Table 1). Most additional phylogenetic studies with macraucheniids in the taxon sampling have either focused on testing relationships within the family (e.g., Püschel et al. 2023) or determining the interordinal relationships within Placentalia (e.g., O’Leary et al. 2013, Buckley 2015) instead of the interfamilial affinities (Table 2).</p><p>Considering the Late Eocene  Polymorphis as the first occurrence of the family and the Pliocene to Late Pleistocene  Macrauchenia (Bond 1999, Prado et al. 2015) and the probably Late Pleistocene  Xenorhinotherium (Cartelle and Lessa 1988) as the last occurrences of  Macraucheniidae, the temporal interval for this family would be around 39.00–~0.11 Mya (Fig. 2B; Supporting information, Table S1, File S2).</p></div>	https://treatment.plazi.org/id/03E487FEFFCAFF85CCDC9A30ACE3276B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFC4FF87CCB49B3CA9F521A5.text	03E487FEFFC4FF87CCB49B3CA9F521A5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Proterotheriidae (Odreman Rivas 1969)	<div><p>Proterotheriidae (Figs 4J–K, 5H–I)</p><p>With 28 currently accepted genera,  Proterotheriidae is the most diverse family of litopterns (Fig. 2A; Supporting information, Table S1), being particularly diverse during the Neogene alongside macraucheniids (Fig. 2B–C). Soria (2001) presented a major revision of this family, so to avoid unnecessary repetition, only the most important taxonomical aspects and the main advances since his revision are discussed here.  Proterotheriidae was named by Ameghino (1887) with the description of the genera  Proterotherium Ameghino, 1883,  Thoatherium Ameghino, 1887,  Diadiaphorus Ameghino, 1887, and  Licaphrium Ameghino, 1887 . Simpson (1945) subdivided this family into two subfamilies,  Polymorphinae and  Proterotheriinae, the former including taxa mostly from the Palaeocene and Eocene (Peligran to Mustersan SALMAs) and the later the taxa from the Late Oligocene onwards (Deseadean SALMA). Although this classification was followed by other authors (Lavocat 1958, Paula Couto 1979), Simpson (1948) did not employ this subfamilial division and only used the family  Proterotheriidae, without giving any explanation for this omission. Later, Odreman-Rivas (1969) modified the subfamily  Polymorphinae from Simpson’s (1945) proposal, keeping  Polymorphis Roth, 1899 (including  Polyacrodon ligatus Roth, 1899 and  Heteroglyphis Roth, 1899), and excluding the rest of the taxa such as the now considered anisolambdids  Wainka Simpson, 1935 and  Anisolambda Ameghino, 1901 (including  Ricardolydekkeria Ameghino, 1901 and  Josepholeidya Ameghino, 1901) and the didolodontid  Xesmodon Berg, 1899 .</p><p>Cifelli(1983a)subdividedthefamilyProterotheriidaeintotwo subfamilies,  Anisolambdinae and  Proterotheriinae, the former subfamily including mostly anisolambdids (see  Anisolambdidae section for more details), and the latter similar to the original definition of Simpson (1945). In addition, Cifelli (1983a) excluded the subfamily  Polymorphinae from  Proterotheriidae, placing it insteadwithinthefamilyMacraucheniidae (seeMacraucheniidae section for more details). Soria (2001) later removed the subfamily  Anisolambdinae from  Proterotheriidae, elevating its rank as a distinct family (see  Anisolambdidae section for more details), which leaves the concept of  Proterotheriidae very similar to the subfamily  Proterotheriinae from Simpson (1945), although with numerous new additions to the family since then (e.g., Corona et al. 2020, McGrath et al. 2020a; Supporting information, Table S1). Important additions to  Proterotheriidae were the bunodont  Megadolodus molariformis McKenna, 1956 and  Neodolodus colombianus Hoffstetter &amp; Soria, 1986 (Cifelli and Diaz 1989, Cifelli and Villarroel 1997, McGrath et al. 2020b), which were previously considered didolodontids (McKenna 1956, Hoffstetter and Soria 1986). Including both taxa within  Litopterna has phylogenetic support, but their affinities within  Proterotheriidae and even other litoptern families are unclear (Carrillo et al. 2023).</p><p>Proterotheriidae has been included by some authors as part of suborder  Lopholipterna (Cifelli 1983a, Soria 2001; Table 1), which also includes the families  Adianthidae and  Macraucheniidae, a hypothesis that later found phylogenetic support (Cifelli 1993; Fig. 1C). Since then, most phylogenetic studies that included proterotheriids have focused either on resolving the affinities within  Proterotheriidae (e.g., McGrath et al. 2020b) or phylogenies studying interordinal relationships between SANUs (e.g., MacPhee et al. 2021) instead of phylogenies examining the interfamilial affinities of proterotheriids (Table 2).</p><p>Following Soria’s (2001) concept of the family, the earliest proterotheriid is  Lambdaconus suinus Ameghino, 1897 from different localities from the Sarmiento Formation, Chubut, Argentina, and the last is  Neolicaphrium recents Frenguelli, 1921 from the Late Pleistocene from different localities in Argentina, Brazil, and Uruguay (Gaudioso et al. 2017), giving a temporal interval of around 29.3–~0.11 Mya to this family (Fig. 2B; Supporting information, Table S1).</p></div>	https://treatment.plazi.org/id/03E487FEFFC4FF87CCB49B3CA9F521A5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFC6FF81CE499C73A8F824B5.text	03E487FEFFC6FF81CE499C73A8F824B5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Adianthidae	<div><p>Adianthidae (Figs 4F–H, 5E–G)</p><p>Adianthidae currently includes six genera, being considerably less diverse than proterotheriids and macraucheniids during the Neogene (Fig. 2; Supporting information, Table S1). Cifelli and Soria (1983a) presented a detailed revision of this family, so to avoid unnecessary repetition, only the most relevant taxonomic aspects are discussed here, along with the main advances since their revision. Known as the pygmy litopterns because of their reduced size, the family  Adianthidae was named by Ameghino (1891a) to place the Early Miocene  Adianthus bucatus Ameghino 1891a from the Santa Cruz Formation, Argentina (Cifelli and Soria 1983a). Among other distinct anatomical features, adianthids present a trilobed m3 and enamel fossettes in the P4–M3 formed by hypertrophied conular cristae (Cifelli and Soria 1983a). Following new fossil discoveries, Ameghino (1906) added three additional genera to the family  Adianthidae:  Proadiantus Ameghino, 1897 and  Tricoelodus Ameghino, 1897 from the Sarmiento Formation (Woodburne et al. 2014a), and  Pseudadiantus Ameghino, 1901 . The last was later excluded from the family and formally synonymized with the notopithecine interatheriid  Antepithecus by Simpson (1967).</p><p>Bordas (1939) added  Proheptaconus Bordas, 1936 from the Colhue-Huapi Member of the Sarmiento Formation (Soria 1981) to what he considered the subfamily  Adianthinae, a subfamily of  Macraucheniidae, which was followed by other authors (Patterson 1940, Simpson and Minoprio 1949). However, Simpson et al. (1962) later restored the family  Adianthidae, but now including a new taxon,  Adiantoides leali Simpson &amp; Minoprio, 1949 from the Divisadero Largo Formation, Argentina (age uncertain; Cerdeño et al. 2008, López 2010, Woodburne et al. 2014a). Bond and Vucetich (1983) added to  Adianthidae Indalecia grandensis Bond &amp; Vucetich, 1983 from the Lumbrera Formation, Argentina (Early Eocene; Fernicola et al. 2021), based on unique shared anatomical features with  Adiantoides leali, such as the absence of a postprotocrista connecting the protocone with the metaconule and a strong parastyle in M1– M2. These features are not present in other adianthids, so they grouped them in the new subfamily  Indaleciinae, keeping both species tentatively within the family  Adianthidae, which was later followed with doubts by Cifelli and Soria (1983a). In the same work, Cifelli and Soria (1983a) added the genus  Thadanius Cifelli &amp; Soria, 1983 from La Salla-Luribay Basin, Bolivia to the family  Adianthidae .</p><p>Cifelli (1983a) revisited the members of the family, and included  Proectocion Ameghino 1904b [previously considered a didolodontid by Simpson (1948)] as the earliest member of  Adianthidae based on dental similarities. In addition, he proposed a new superfamily called Macrauchenioidea, to group adianthids and macraucheniids based on dental similarities, such as an absent paraconid and the presence of a paralophid in the lower molars (Table 1). Soria (1984a), based on dental similarities with ‘amilnedwarsids’, elevated the rank of the subfamily  Indaleciinae to the family  Indaleciidae, and later Soria (1989b) classified them as a family of the order  Notopterna (see sections of  Amilnedwardsiidae and the order  Notopterna, and the family  Indaleciidae).  Adianthidae has also been considered by some authors as part of the suborder  Lopholipterna (Cifelli 1983a, Soria 2001), which includes the families  Macraucheniidae and  Proterotheriidae (Table 1), a hypothesis that later found phylogenetic support (Cifelli 1993; Fig. 1C). In the same study, Macrauchenioidea also found support, but only when indaleciids are excluded (Cifelli 1993). Strikingly, there has been not a single study examining the phylogenetic relationships within  Adianthidae using modern phylogenetic methods, and most recent phylogenetic studies have included adianthids as outgroups for phylogenies of  Macraucheniidae (e.g., Forasiepi et al. 2016; Table 2).</p><p>If we follow Cifelli (1983a) and consider  Proectocion from the Barranca south of Lago Colhue-Huapi of Argentina (Middle Eocene; Kramarz and Bond 2013, Woodburne et al. 2014a) as the earliest member of  Adianthidae and considering  Adianthus godoyi from the Río Frías Formation at Alto Río Cisnes, Chile as the last member of this family (Cifelli 1991, de la Cruz and Cortés 2011), the family  Adianthidae encompasses a temporal interval of 42–14.8 Mya (Fig. 2B; Supporting information, Table S1).</p></div>	https://treatment.plazi.org/id/03E487FEFFC6FF81CE499C73A8F824B5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFC0FF83CE7D9B62AA2B26A2.text	03E487FEFFC0FF83CE7D9B62AA2B26A2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sparnotheriodontidae	<div><p>Sparnotheriodontidae (Figs 4L, 5L)</p><p>Sparnotheriodontidae currently has five accepted genera, all with a fossil record exclusively in the Palaeogene (Fig. 2; Supporting information, Table S1). The family  Sparnotheriodontidae was initially proposed by Soria (1980a) to include  Sparnotheriodon epsilonoides Soria, 1980a from Cañadón Vaca Member, Sarmiento Formation, Argentina, which was classified as incertae sedis within the order  Notoungulata . The family was later removed from  Notoungulata and incorporated into the order  Litopterna, being expanded by Soria (1980b) adding the genera  Victorlemoinea and  Phoradiadus Simpson et al. 1962 .  Victorlemoinea was previously considered an early macraucheniid (Simpson 1945, 1948), and  Phoradiadus was previously considered with doubts as a proterotheriid (Simpson et al. 1962). Soria’s (1980b) proposal was based on shared similarities in the upper and lower molars, such as the presence of a lophoid metaconule and a closed trigon basin in M2.  Phoradiadus includes one species,  Phoradiadus divortiensis, from Divisadero Largo Formation, Mendoza, Argentina (age uncertain; Cerdeño et al. 2008, López 2010, Woodburne et al. 2014a). An interesting feature of the teeth of the  Sparnotheriodontidae is that they possess vertical Hunter–Schreger bands, a feature unknown in any other litoptern (Bond et al. 2006).</p><p>Simpson (1948) tentatively accepted three Patagonian species of the genus  Victorlemoinea:  Victorlemoinea labyrinthica Ameghino, 1901,  Victorlemoinea emarginata Ameghino, 1901, and  Victorlemoinea longidens Ameghino, 1901, as they were represented only by isolated teeth, the first two being represented by upper teeth, and the last by lower teeth. This means that  V. longidens could correspond to teeth from the lower dentition of either  V. labyrinthica or  V. emarginata, in particular the mandibular fragment with m1–m2 of the holotype (MACN A-10670), as the allegedly associated premolars of this specimen seem to correspond to an isotemnid notoungulate (Bond et al. 2006).  In the genus  Victorlemoinea, Bond et al. (2006) included only  V. labyrinthica in their list of formally recognized sparnotheriodontid species, and more recently some authors have mentioned only  V. labyrinthica for the  Riochican and  Vacan Patagonian faunas, omitting  V. emarginata and  V. longidens (e.g., Reguero et al. 2014, Gelfo 2016, Gelfo et al. 2019), which can be interpreted as implicitly synonymizing them with  V. labyrinthica . In addition, there is one  Victorlemoinea species from Itaboraí, Brazil,  Victorlemoinea prototypica Paula Couto, 1952, also known mostly from dental material.</p><p>Cifelli (1983b) indirectly assigned some tarsals (i.e., calcaneum and astragalus) to  V. prototypica, based on their relative size and abundance. These tarsals were large and scarce, so considering linear regressions with the dentition (m2 area), he argued it could only correspond to either the sparnotheriodontid  V. prototypica or the didolodontid  Lamegoia conodonta . Considering the similarities of the tarsals assigned to  V. prototypica and others indirectly assigned to didolodontids, such as presenting a medial malleolar facet of the astragalus extending onto the neck and the presence of a dorsal beak on the distal end of the calcaneum,Cifelli (1983a) grouped  Sparnotheriodontidae and  Didolodontidae under the superfamily  Didolodontoidea in the order  Condylarthra (Table 1). This hypothesis found some phylogenetic support (Cifelli 1993, Bergqvist 1996; Fig. 1C), and has been followed by some authors (e.g., Bergqvist 2008). However, other authors have questioned the indirect anatomical assignment of these tarsals to  V. prototypica (e.g., Hoffstetter and Soria 1986, Soria 2001, Lorente 2015), and even suggested an affinity of these tarsals with  Notoungulata (Soria 2001) or with  Astrapotheria (Lorente 2015) .</p><p>Among the different litoptern families, Soria (2001) considered that sparnotheriodontids were more closely related to the family  Anisolambdidae due to overall dental similarities between both families, and also considering an isolated M1 (MNRJ 1479- V) from Itaboraí showing an intermediate anatomy between sparnotheriodontids and anisolambdids. Therefore, he created the suborder  Eolitopterna to include both families (Table 1). In addition, Soria (2001) tentatively added  Heteroglyphis dewoletzky Roth, 1899 to  Sparnotheriodontidae without any detailed anatomical justification; this species is known from a broken upper molar probably from Cerro del Humo, Argentina. This taxon was previously referred to  Proterotheriidae by Simpson (1948).</p><p>More recently, one genus and two new species from the Eocene La Meseta and Submeseta formations, Antarctica, have been added to the family  Sparnotheriodontidae,  Notiolofos arquinotiensis Bond et al. 2006 and  Notiolofos regueroi Gelfo, López &amp; Santillana, 2017 . They are differentiated mainly by their size and represented by isolated teeth (the latter species only by a m3; Gelfo et al. 2019).  Notiolofos arquinotiensis is particularly interesting as it is the most abundant land mammal in</p></div>	https://treatment.plazi.org/id/03E487FEFFC0FF83CE7D9B62AA2B26A2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFC3FF82CF82999BA9FC2368.text	03E487FEFFC3FF82CF82999BA9FC2368.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Amilnedwardsiidae	<div><p>Amilnedwardsiidae and the order  Notopterna</p><p>Amilnedwardsiidae was proposed preliminarily by Soria (1984b) and better justified in a later article (Soria 1989a). It is based on two genera (i.e., Amilnedwarsia Ameghino, 1901 and  Rutimeyeria Ameghino, 1901) and three species named after small bunodont isolated upper molars whose positions were assigned tentatively by Soria (1989a) by comparing them with the indaleciid  Indalecia grandensis .  Amilnedwardsiidae was considered to be the most basal family of the order  Notopterna Soria, 1989, which also includes the families  Indaleciidae and  Notonychopidae (Soria 1989b) .  Notopterna is diagnosed by a well-developed and buccally projected parastyle and the absence of a mesostyle in the upper molars, bicrescentic and selenodont lower molars, and anteriorly acuminate petrosals, among other anatomical features that distinguish them from other SANUs (Soria 1989a). Soria (1989a, b) argued that, considering their dental similarities, ‘amilnedwarsids’ could be the structural ancestor of both indaleciids and notonychopids. However, a phylogenetic analysis failed to recover the members of  Notopterna as a monophyletic group (Bonaparte and Morales 1997), which has led to some authors to include its members tentatively within  Litopterna (e.g., Gelfo et al. 2016, Croft et al. 2020, Goin et al. 2022). However, this is a problematic assumption as Bonaparte and Morales (1997) used a matrix that includes only litopterns, so the broader relationships of notopterns have not yet been tested in an analysis that includes other SANUs such as notoungulates and astrapotheres.</p><p>Amilnedwardia and  Rutimeyeria were previously referred tentatively to the family  Macraucheniidae and regarded as close to  Victorlemoinea (Simpson 1945, 1948), although the latter now is now considered to be part of the family  Sparnotheriodontidae . Cifelli (1983a)considered referred specimens of these taxa to be too fragmentary to determine a clear affinity,but suggested the referred specimens represent deciduous teeth of a henricosborniid notoungulate based on the presence of a metaconule-derived crest connecting with the ectoloph wall. Indeed, probably due to their limited fossil record, thus far no members of this family have ever been included in a phylogenetic analysis.</p><p>If we consider  Amilnedwardsiidae to be a valid family, as its three species all come from Patagonian fossil localities of a similar age that correlate with the Vacan SALMA, the temporal interval of this family is ~45–42 Mya (Fig. 2B; Supporting information, Table S1; Krause et al. 2017).</p></div>	https://treatment.plazi.org/id/03E487FEFFC3FF82CF82999BA9FC2368	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFC3FF9CCE5B9F05AA13276B.text	03E487FEFFC3FF9CCE5B9F05AA13276B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Indaleciidae (Bond & Vucetich 1983)	<div><p>Indaleciidae (Figs 6A, 7A)</p><p>Indaleciidae currently includes three accepted genera with an exclusive Palaeogene distribution (Fig. 2; Supporting information, Table S1).  Indaleciidae was initially proposed by Soria (1984a) and better justified in a later article (Soria 1989a), in which he elevated the rank of the previously proposed subfamily  Indaleciinae, which included  Indalecia grandensis and  Adiantoides leali (Bond and Vucetich 1983) . Previously,  A. leali from Divisadero Largo Formation, Argentina (age uncertain; Cerdeño et al. 2008, Woodburne et al. 2014a, López 2010), was placed tentatively in  Adianthidae based on dental similarities with the adianthid  Adianthus (Simpson and Minoprio 1949, Simpson et al. 1962). Later, Bond and Vucetich (1983) described  Indalecia grandensis from the Lumbrera Formation, Argentina [Early Eocene (Fernicola et al. 2021)], that showed dental similarities with  Adiantoides leali, such as a buccolingually transverse crest connecting the hypocone with the metaconule, and the absence of a postprotocrista connecting the protocone with the metaconule on M1–M2, among other features that justified the proposal of a subfamily  Indaleciinae within  Adianthidae . Cifelli and Soria (1983a) tentatively followed Bond and Vucetich (1983) considering the shared presence of fossettes formed by hypertrophied conular cristae in the upper molars in members of  Indaleciinae and typical adianthids. However, they also noticed some important features that separated them, such as the lack of a mesostyle on the upper molars and the lack of an entolophid (hypolophid in our matrix nomenclature; Fig. 3) on the lower molars, among other features. Cifelli and Soria (1983a) also added a new species  Adiantoides magnus from Cañadon Vaca, Argentina, to  Indaleciinae . The differences between indaleciines and adianthids and also the dental similarities of the former with ‘amilnedwarsiids’, led Soria (1989a) to propose elevating the subfamily  Indaleciinae to the family  Indaleciidae, including indaleciids, ‘amilnedwarsiids’, and notonychopids under the same order  Notopterna (see  Amilnedwardsidae and the order  Notopterna section). Apart from some common dental features, indaleciids share features of the ear region with notonychopids, such as anteriorly acuminate petrosals; however, they diverge in the presence of enamel fossettes in the P2– M3 and less developed parastyles in indaleciids, among other dental features (Soria 1989b).</p><p>Cifelli (1993) conducted a phylogenetic analysis that included indaleciids (i.e.,  Indalecia and  Adiantoides) in a sample of different litoptern families and didolodontids, and found them to form a monophyletic group closely related to sparnotheriodontids (i.e.,  Victorlemoinea), and these three taxa together were sister to didolodontids (Fig. 1C). This result was probably influenced by Cifelli’s (1983b) indirect assignment of tarsals for  Victorlemoinea (see  Sparnotheriodontidae section). When Bonaparte and Morales (1997) repeated the analysis removing didolodontids and adding notonychopids, they similarly found this close relationship between indaleciids and sparnotheriodontids, without a close relationship with notonychopids (Fig. 1D). However, in a different analysis based only on lower molars, Gelfo et al. (2008) found indaleciids in a polytomy with  Notonychops powelli, and this polytomy as sister of the sparnotheriodontid  Sparnotheriodon . Most authors since then have included  Indaleciidae as a family of  Litopterna (e.g., Croft et al. 2020). Nevertheless, a preliminary study that included ear region characters among other craniodental characters found the indaleciid  Indalecia as a sister taxon to astrapotheres and notoungulates, instead of having a close relationship with  Litopterna (García-López and Babot 2014) . More recently, in a study that included litopterns, kollpanines, and didolodontids, the indaleciid  Adiantoides was found at the stem of litopterns, being more closely related to them than to didolodontids (Kramarz et al. 2021). In a different study that also included some early notoungulates, the indaleciid  Indalecia was found in a polytomy that includes sparnotheriodontids, protolipternids, didolodontids, and North American phenacodontids (Zimicz et al. 2022). It is important to mention that most of these studies have a relatively limited taxon and character sample, and none of them have included members of all the litoptern families and early members of the various SANU orders (Table 2). Therefore, including  Indaleciidae within the order  Litopterna can only be considered as tentative.</p><p>The earliest member of  Indaleciidae is the Early Eocene  Indalecia grandensis from the Lumbrera Formation, Argentina (55.0–46.2 Mya; Fernicola et al. 2021) and the youngest would be an undescribed indaleciid from the Abanico Formation, Chile (Wyss et al. 1994), which gives the family a temporal interval of 55.0–~31.5 Mya (Fig. 2B; Supporting information, Table S1; Flynn et al. 2003, Fernicola et al. 2021).</p></div>	https://treatment.plazi.org/id/03E487FEFFC3FF9CCE5B9F05AA13276B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFDDFF9CCE779B3CAD3A264F.text	03E487FEFFDDFF9CCE779B3CAD3A264F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Notonychopidae (Soria 1989)	<div><p>Notonychopidae (Figs 6B–C, 7B)</p><p>Notonychopidae currently includes two accepted genera with an exclusively Palaeogene distribution (Fig. 2; Supporting information, Table S1).  Notonychopidae was founded to include  Notonychops powelli Soria, 1989 from the Rio Loro Formation [age uncertain, although probably pre-Itaboraian SALMA age (Gelfo et al. 2020b)], an ungulate with great development of the parastyle in M1–M3, which among other features, makes it anatomically distinctive from any other SANU previously described (Soria 1989b). Soria (1989b) considered that  N. powelli was dentally convergent with members of the family  Esthonychidae (Tillodonta), particularly with the lower molars being brachydont and selenodont with a subequal trigonid and talonid length. However,  N. powelli shares important anatomical features with the dentition of ‘amilnedwarsids’ and indaleciids, and of the ear region with indaleciids, which madeSoria (1989b) place it within the order  Notopterna (see  Amilnedwardsidae and the order  Notopterna section for more details). Key differences between notonychopids with indaleciids are the absence of enamel fossettes in the P2–M3 and a greater development of the parastyle in the former, among other features (Soria 1989b). McKenna and Bell (1997) later classified  Notonychopidae as a family within the order  Litopterna, albeit without any anatomical justification (Table 1). Bonaparte and Morales (1997) added to  Notonychopidae the taxon  Requisia vidmari Bonaparte and Morales 1997 from Punta Peligro (Peligran SALMA) which they found to be phylogenetically close to  Wainka tshotshe (Fig. 1D). However, since this analysis, Goin et al. (2022) called into question an isolated m3 (UNP 946) previously assigned to  Requisia vidmari by Bonaparte and Morales (1997) because of its bunodont features and how low and different its cristids are to what would be expected for a notonychopid, suggesting it could be a member of  Didolodontidae .</p><p>Bonaparte and Morales (1997) failed to find a monophyletic  Notonychopidae, as  N. powelli and  R. vidmari were not as closely related in their phylogeny (Fig. 1D). In addition, as notonychopids were found nested within  Litopterna, most authors now consider them litopterns (e.g., Gelfo et al. 2016, Croft et al. 2020, Goin et al. 2022), which is problematic given that Bonaparte and Morales (1997) used a matrix that includes exclusively litopterns and did not test alternative relationships. Since the early phylogenetic study of Bonaparte and Morales (1997), there has not been any attempt to test the monophyly of  Notonychopidae and to resolve its position among litopterns and other SANUs (Table 2).</p><p>As the only two known members of  Notonychopidae are  R. vidmari and the potentially pre-Itaboraian SALMA  N. powelli, the temporal interval of this family is not well constrained with a current range of ~63.8–51.4 Mya, and is likely much narrower in range (Fig. 2B; Supporting information, Table S1; Woodburne et al. 2014b, Krause et al. 2017, Gelfo et al. 2020b).</p></div>	https://treatment.plazi.org/id/03E487FEFFDDFF9CCE779B3CAD3A264F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
03E487FEFFDDFF98CCA29A59A919272A.text	03E487FEFFDDFF98CCA29A59A919272A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Didolodontidae	<div><p>Didolodontidae (Figs 8B–G, 9B–E)</p><p>Didolodontidae currently includes 10 genera [or 12 if we include two unnamed genera (Tejedor et al. 2009)] with an exclusive Palaeogene distribution (Fig. 2B; Supporting information, Table S1). There has been recent major detailed revision of this family (Gelfo 2006, 2010), so to avoid unnecessary repetition, only the most relevant taxonomic aspects are discussed here, along with the advances since these revisions. The family  Didolodontidae was proposed by Scott (1913) to classify a group of bunodont ‘condylarths’ (see ‘Condylarthra’ section in Supporting information, File S1) endemic to South America with a Palaeogene fossil record consisting mostly of isolated teeth. Initially, Ameghino (1897) described the first known Eocene didolodontid species  Didolodus multicuspis as a phenacodontid. Osborn (1910) identified additional South American taxa as ‘Condylarthra’ incertae sedis such as  Notoprotogonia (Ameghino 1904b [later synonymized with  Ernestokokenia Ameghino, 1901 by Simpson (1948)],  Lambdaconus [later considered an early proterotheriid (Soria 2001)], and  Proectocion [later considered an early adianthid (Cifelli 1983a)]. Scott (1913) later grouped these four taxa as part of the family  Didolodidae within the order  Litopterna (Table 1). Simpson (1934) subsequently corrected it to  Didolodontidae, and grouped them again within</p><p>‘Condylarthra’. After adding two new species from the Rio Chico Formation [latest Danian–Late Lutetian in age (Krause et al. 2017)] of the genus  Ernestokokenia ( Ernestokokenia yirunhor Simpson, 1935 and  Ernestokokenia chaishoer Simpson 1935), Simpson (1948) revised the whole group, giving a new definition for  Didolodontidae and emending many of Ameghino’s (1906) taxonomic classifications of the Patagonian taxa, including the following genera in the family:  Didolodus,  Ernestokokenia,  Argyrolambda Ameghino 1904b,  Paulogervaisia Ameghino, 1901,  Proectocion,  Enneoconus Ameghino, 1901, Asmithwoodwarsia Ameghino, 1901, and  Archaeohyracotherium Ameghino, 1906 . Simpson soon recognized the close similarities between this group and North American ‘condylarths’, including species of  Phenacodontidae and  Mioclaenidae, stating that they were impossible to separate on a purely morphological basis (Simpson 1948, 1980).</p><p>Paula Couto (1952) added two species of  Ernestokokenia ( Ernestokokenia protocenica Paula Couto, 1952 and  Ernestokokenia parayiruhnor Paula Couto, 1952) and the new genus  Lamegoia from fissure fills in the Itaboraí Formation, Brazil, to the family  Didolodontidae . McKenna (1956) added  Megadolodus molariformis from La Venta, Colombia, which Cifelli and Villarroel (1997) later considered to be a bunodont proterotheriid, a hypothesis that recently has found some support in phylogenetic analyses (Carrillo et al. 2023). Paula Couto (1978) added  Asmithwoodwardia scotti Paula Couto, 1952 from Itaboraí to the family, a taxon that he previously classified within the North American ‘Condylarthra’ family  Hyopsodontidae based on dental similarities (Paula Couto 1952). In a revision of the family  Didolodontidae, Cifelli (1983a), based on an indirect tarsal association, removed  Ernestokokenia parayirunhor (renamed as  Miguelsoria parayirunhor Cifelli 1983a) and  Asmithwoodwardia scotti from  Didolodontidae and placed them in  Litopterna (see  Protolipternidae section for more information). This tarsal association was based on the fact that the relative size and abundance of these tarsals in the Itaboraí sites was what you would expect for the size and abundance of the dentition of  Miguelsoria parayirunhor (Cifelli 1983b) . Cifelli (1983a) reclassified  Ernestokokenia protocenica as  Paulacoutoia protocenica [now known as  Ricardocifellia protocenica (Mones, 2015) because of preoccupation], because he considered it sufficiently morphologically distinct from the genus  Ernestokokenia . According to Cifelli (1983a), didolodontids are characterized by having a primitive dentition very similar to mioclaenids, but possessing a hypocone on the M3 (absent in mioclaenids) and a primitive ankle structure similar to many South and North American ‘condylarths’ (i.e., the astragalar facet for the medial malleolus is enlarged anteriorly and the calcaneal cuboid facet is dorsally expanded). Soria (2001), based on dental features, removed  Xesmodon Berg, 1899 from  Anisolambdidae and placed it within  Didolodontidae .</p><p>In a detailed revision of  Didolodontidae, Gelfo (2006) synonymized  Argyrolambda with  Didolodus,  Archaeohyracotherium with  Asmithwoodwardia, and  Enneoconus with  Ernestokokenia, thus reducing the total number of valid didolodontid genera. Later, Gelfo (2007), based on dental similarities and a phylogenetic analysis, added  Escribania chubutensis Bonaparte et al., 1993 and  Raulvaccia peligrensis Bonaparte et al. 1993 from Punta Peligro, Argentina, to  Didolodontidae .  Escribania chubutensis and  Raulvaccia peligrensis were previously classified within the ‘condylarth’ North American family  Mioclaenidae (Bonaparte et al. 1993) . The most recent generic additions to  Didolodontidae have been  Umayodus Gelfo &amp; Sigé, 2011 from the Lower Muñani Formation, Peru, and  Saltaodus Gelfo et al., 2020a from the Lumbrera Formation, both of uncertain age (Gelfo and Sigé 2011, Zimicz et al. 2023).</p><p>In terms of the phylogenetic affinities of  Didolodontidae, early phylogenetic analyses found a close relationship between didolodontids and sparnotheriodontids (Cifelli 1993, Bergqvist 1996; Fig. 1C), but these were based on tarsal associations that have been questioned by later authors (e.g.,Soria 2001, Gelfo and Sigé 2011; see also  Sparnotheriodontidae and  Protolipternidae sections). Muizon and Cifelli (2000), in a phylogenetic analysis that included North American mioclaenids, South American kollpaniines, didolodontids, and protolipternids, found that protolipternids were nested within  Didolodontidae and that both groups together formed the sister group of kollpanines, a result that was also found in a later study (Gelfo 2006). This means that  Didolodontidae excluding  Protolipternidae is a paraphyletic group. Other phylogenetic studies that have included didolodontids in very small matrices have focused on their internal relationships, assuming their monophyly (e.g., Gelfo 2010; Table 2).</p><p>When didolodontids have been included with a limited taxon sample (i.e., usually  Didolodus) in broader analyses that include other SANUs, they have been found in a branch at the stem of some North American ‘condylarths’ and other SANUs (e.g., Billet et al. 2015), or in a basal position in Laurasiatheria as the sister group of  Protolipternidae (O’Leary et al. 2013) . When a larger sample of didolodontids has been included, they have been found at the stem of litopterns but not forming a monophyletic  Didolodontidae (Kramarz et al. 2021), or as a paraphyletic group among SANUs and North American ‘condylarths’ (Zimicz et al. 2023). In sum, there is still a considerable uncertainty in the phylogenetic position of didolodontids and their relationship among SANUs, and whether they form a monophyletic group. In fact, there has not been any phylogenetic analysis that includes didolodontids alongside representatives of all litoptern families, protolipternids, and the different SANUs’</p></div>	https://treatment.plazi.org/id/03E487FEFFDDFF98CCA29A59A919272A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Püschel, Hans P.;Shelley, Sarah L.;Williamson, Thomas E.;Perini, Fernando A.;Wible, John R.;Brusatte, Stephen L.	Püschel, Hans P., Shelley, Sarah L., Williamson, Thomas E., Perini, Fernando A., Wible, John R., Brusatte, Stephen L. (2024): A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates. Zoological Journal of the Linnean Society 202 (1): 1-50, DOI: 10.1093/zoolinnean/zlae095, URL: http://dx.doi.org/10.1093/zoolinnean/zlae095
